Biol. Rev. (2010), 85, pp. 55–110.

55
doi:10.1111/j.1469-185X.2009.00094.x

The origin and early evolution of dinosaurs

Max C. Langer1∗ , Martin D. Ezcurra2 , Jonathas S. Bittencourt1
and Fernando E. Novas2,3
1 Departamento de Biologia, FFCLRP, Universidade de São Paulo; Av. Bandeirantes 3900, Ribeirão Preto-SP, Brazil
2
Laboratorio de Anatomia Comparada y Evolución de los Vertebrados, Museo Argentino de Ciencias Naturales ‘‘Bernardino Rivadavia’’, Avda.
Angel Gallardo 470, Cdad. de Buenos Aires, Argentina
3 CONICET (Consejo Nacional de Investigaciones Cient ı́ficas y Técnicas); Avda. Rivadavia 1917 - Cdad. de Buenos Aires, Argentina

(Received 28 November 2008; revised 09 July 2009; accepted 14 July 2009)

ABSTRACT

The oldest unequivocal records of Dinosauria were unearthed from Late Triassic rocks (approximately 230 Ma)
accumulated over extensional rift basins in southwestern Pangea. The better known of these are Herrerasaurus
ischigualastensis, Pisanosaurus mertii, Eoraptor lunensis, and Panphagia protos from the Ischigualasto Formation, Argentina,
and Staurikosaurus pricei and Saturnalia tupiniquim from the Santa Maria Formation, Brazil. No uncontroversial
dinosaur body fossils are known from older strata, but the Middle Triassic origin of the lineage may be inferred
from both the footprint record and its sister-group relation to Ladinian basal dinosauromorphs. These include
the typical Marasuchus lilloensis, more basal forms such as Lagerpeton and Dromomeron, as well as silesaurids: a possibly
monophyletic group composed of Mid-Late Triassic forms that may represent immediate sister taxa to dinosaurs.
The first phylogenetic definition to fit the current understanding of Dinosauria as a node-based taxon solely
composed of mutually exclusive Saurischia and Ornithischia was given as ‘‘all descendants of the most recent
common ancestor of birds and Triceratops’’. Recent cladistic analyses of early dinosaurs agree that Pisanosaurus mertii
is a basal ornithischian; that Herrerasaurus ischigualastensis and Staurikosaurus pricei belong in a monophyletic Her-
rerasauridae; that herrerasaurids, Eoraptor lunensis, and Guaibasaurus candelariensis are saurischians; that Saurischia
includes two main groups, Sauropodomorpha and Theropoda; and that Saturnalia tupiniquim is a basal member of
the sauropodomorph lineage. On the contrary, several aspects of basal dinosaur phylogeny remain controversial,
including the position of herrerasaurids, E. lunensis, and G. candelariensis as basal theropods or basal saurischians,
and the affinity and/or validity of more fragmentary taxa such as Agnosphitys cromhallensis, Alwalkeria maleriensis,
Chindesaurus bryansmalli, Saltopus elginensis, and Spondylosoma absconditum. The identification of dinosaur apomorphies
is jeopardized by the incompleteness of skeletal remains attributed to most basal dinosauromorphs, the skulls and
forelimbs of which are particularly poorly known. Nonetheless, Dinosauria can be diagnosed by a suite of derived
traits, most of which are related to the anatomy of the pelvic girdle and limb. Some of these are connected to the
acquisition of a fully erect bipedal gait, which has been traditionally suggested to represent a key adaptation that
allowed, or even promoted, dinosaur radiation during Late Triassic times. Yet, contrary to the classical ‘‘compet-
itive’’ models, dinosaurs did not gradually replace other terrestrial tetrapods over the Late Triassic. In fact, the
radiation of the group comprises at least three landmark moments, separated by controversial (Carnian-Norian,
Triassic-Jurassic) extinction events. These are mainly characterized by early diversification in Carnian times, a
Norian increase in diversity and (especially) abundance, and the occupation of new niches from the Early Jurassic
onwards. Dinosaurs arose from fully bipedal ancestors, the diet of which may have been carnivorous or omnivo-
rous. Whereas the oldest dinosaurs were geographically restricted to south Pangea, including rare ornithischians
and more abundant basal members of the saurischian lineage, the group achieved a nearly global distribution by
the latest Triassic, especially with the radiation of saurischian groups such as ‘‘prosauropods’’ and coelophysoids.

Key words: Dinosauria, Dinosauromorpha, Triassic, phylogeny, evolution, biogeography, Herrerasauria.

∗Address for correspondence: Departamento de Biologia, Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto, Universidade
de São Paulo; Av. Bandeirantes 3900, 14040-901, Ribeirão Preto-SP, Brazil (Tel: +55 16 3602 3844; Fax: +55 16 3602 4886;
E-mail: mclanger@ffclrp.usp.br)

Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
56 Max C. Langer and others

CONTENTS
I. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
(1) Historical background on early dinosaurs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
(2) The dinosauromorph radiation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
II. Phylogeny and Systematics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
(1) What makes a dinosaur? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
(2) Phylogenetic definitions: naming early dinosaurs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
III. Dinosaur ‘‘Trail Blazers’’ in Space, Time, and Evolutionary Context . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
(1) The oldest dinosaurs and the rocks that contain them . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
(2) The evolutionary tree of early dinosaurs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
(3) Geographical distribution of basal dinosaurs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
IV. Ecology of the Dinosaur Radiation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
(1) The Triassic scene . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
(2) Lucky break? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
(3) Of legs and teeth: insights on the palaeobiology of early dinosaurs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
V. Outcomes of a Radiation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
(1) Early ornithischian evolution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
(2) Early sauropodomorph evolution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
(3) Early theropod evolution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
VI. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
VII. Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
VIII. Appendix 1. Institutional Abbreviations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
IX. References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94

nested within Theropoda, or regarded as non-eusaurischan

I. INTRODUCTION
Dinosaurs originated in the Triassic period, and the
Late Triassic represents the first 30 of the 165 million
years of their ‘‘non-avian’’ history on Earth. Yet, of
the 500-700 ‘‘well established’’ dinosaur genera (Wang
& Dodson, 2006; Olshevsky, 2007), only about 30
(approximately 5%) were excavated from Triassic rocks,
and the diversity/diversification of the group is mainly
concentrated in the Jurassic (Rauhut, 2005b; Lloyd et al.,
2008) and/or Cretaceous (Wang & Dodson, 2006) periods.
This is especially the case if one accepts the inference of
Wang & Dodson (2006) that the Late Triassic represents
the best sampled subperiod of the entire Mesozoic in terms
of documented dinosaur diversity. Indeed, dinosaurs are
rare in most Triassic fossil assemblages in which they occur,
although by the end of the period they were already dominant
members of various palaeocommunities.
Triassic dinosaurs were mostly bipedal, and not exception-
ally large. The basal-most forms were probably omnivorous,
but predatory and probably herbivorous dinosaurs also
occurred during Late Triassic times. This includes Her-
rerasaurus ischigualastensis, a top predator up to 4 m long
(Novas, 1997a), and Riojasaurus incertus, a plant-eater of about
three tons (Seebacher, 2001). In taxonomic terms, most Tri-
assic dinosaurs are regarded as members of one of the three
major lineages of the group: Theropoda, Sauropodomorpha,
and Ornithischia. Yet, despite representing well-known taxa,
other Triassic dinosaurs have a debated phylogenetic posi-
tion. This is particularly the case of the herrerasaurs, which
were placed basal to the Ornithischia-Saurischia dichotomy,
saurischians.
Appealing inferences on dinosaur palaeobiology can
be drawn from eggs and nestlings (Bonaparte & Vince,
1979; Moratalla & Powell, 1994), monospecific assemblages
(Coombs, 1990; Schwartz & Gillette, 1994), visual-display-
related morphological features (Vickaryous & Ryan, 1997),
and ‘‘stomach contents’’ (Novas, 1997a; Nesbitt et al., 2006)
of Triassic dinosaurs. Yet, the most debated aspect of early
dinosaur macroevolution corresponds to their first radiation,
and various scenarios were invoked to explain the rise of
the clade in a time interval during which most terrestrial
tetrapods suffered important diversity losses. In fact, by Late
Triassic times, dinosaurs arose and took their first steps
along the evolutionary road, and the investigation of their
obscure origins is crucial for the understanding of dinosaur
interrelationships and palaeobiology as a whole.
(1) Historical background on early dinosaurs
Research on early dinosaurs can be said to have started
with the work of the German palaeontologist Friedrich von
Huene, and his descriptions of Saltopus elginensis Huene,
1910 (Fig. 1A), and Spondylosoma absconditum Huene, 1942.
These two forms have completely different provenances,
coming respectively from the Elgin area, in Northern
Scotland, and Rio Grande do Sul, in South Brazil, but
share curious similarities. Both were regarded as saurischian
dinosaurs by Huene (1910, 1942) and were found in deposits
considered the oldest dinosaur-bearing rocks known at the
time. Huene (1932, 1942) identified various other putative
Triassic dinosaurs as equivalent in age to either Saltopus or
Spondylosoma, but most of these were shown to have doubtful

Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
The origin and early evolution of dinosaurs 57

dinosaur affinities (Galton, 1985b; Benton, 1986b; Galton &
Walker, 1996; Benton et al., 2000; Rauhut & Hungerbühler,
2000; Parker et al., 2005; Nesbitt, Irmis & Parker, 2007).
Notable exceptions are Thecodontosaurus antiquus (Benton et al.,
2000) and the material Cope (1889) originally assigned to
Coelophysis bauri (Nesbitt et al., 2007), but these came from
strata currently considered younger (Benton et al., 2000;
Langer, 2005b; Nesbitt et al., 2007). Indeed, the older age of
both the ‘‘Stagonolepis-beds’’ of Elgin (Huene, 1908) and the
‘‘Rio do Rasto’’ [sic] beds at Chiniquá (Huene & Stahlecker,
1931) was corroborated by recent work. The Lossiemouth
Sandstone Formation has been dated as Carnian (Benton &
Walker, 1985), whereas the Dinodontosaurus Assemblage-Zone
of the Santa Maria Formation is considered of Ladinian
age (Langer et al., 2007c); or early-middle Carnian, following
recent modifications on the Late Triassic time-scale (Muttoni
et al., 2004) and the corrections on the radiometric dating of
the Ischigualasto Formation (Furin et al., 2006).
Although the ages of the Lossiemouth Sandstone and
Santa Maria formations were more securely established,
the dinosaur affinities of Saltopus elginensis and Spondylosoma
absconditum are still debated (Rauhut & Hungerbühler, 2000;
Galton, 2000; Langer, 2004). This is in part due to the
poor preservation of the specimens, which do not allow a
comprehensive assessment of their morphological features.
Therefore, it was not until Reig (1963) placed Herrerasaurus
ischigualastensis (Fig. 1B) and Ischisaurus cattoi within Saurischia
that unequivocal early dinosaurs were known to science. The
described specimens were collected in 1961 from deposits
of the Ischigualasto Formation, San Juan province, north-
western Argentina, which have yielded remains attributable
to dinosaurs since the late 1950s (Reig, 1963). With the
discovery, in 1962, of the ornithischian Pisanosaurus mertii
Casamiquela, 1967 (Fig. 1C), in that same stratigraphic unit,
the presence of both main dinosaur lineages (i.e. Ornithischia
and Saurischia), in the Triassic of South America was
confirmed. Another important ‘‘early dinosaur’’ study of
the time was the description of Staurikosaurus pricei Colbert,
1970 (Fig. 1D). Its type and only specimen, discovered in
1936 in the Santa Maria beds of South Brazil, was the first
consensual early dinosaur to be collected.
While the 1980s were quiet times regarding the study
of early dinosaurs, mainly witnessing the description of
incomplete specimens (Galton, 1985b, 1986; Novas, 1986;
Chatterjee, 1987; Murry & Long, 1989), the early nineties
came with new and exciting discoveries. These include the
unearthing, also from the Ischigualasto Formation, of a new
basal dinosaur still to be fully described, Eoraptor lunensis
(Sereno et al., 1993; Sereno, 2007b), and of further material
of Herrerasaurus ischigualastensis (Sereno & Novas, 1992, 1993;
Novas, 1993; Sereno, 1993). In the late nineties, a new series
of discoveries in Rio Grande do Sul, South Brazil, enlarged
the knowledge of early dinosaur diversity. The then basal-
most member of the sauropodomorph lineage, Saturnalia
tupiniquim (Langer et al., 1999; Langer, França & Gabriel,
2007b; Langer, 2003), was unearthed from the Hyperodapedon
Assemblage-Zone of the Santa Maria Formation, whereas
the overlying Caturrita Formation yielded the saurischian
Guaibasaurus candelariensis (Bonaparte, Ferigolo & Ribeiro,
1999; Bonaparte et al., 2007). Since the beginning of this
century, some putative basal dinosaurs have been described
(Fraser et al., 2002; Ferigolo & Langer, 2007; Nesbitt et al.,
2007; Nesbitt & Chatterjee, 2008; Martinez & Alcober, 2009;
Ezcurra, 2008), while the validity of others was evaluated in
the light of new evidence (Remes & Rauhut, 2005; Yates,
2007b). More importantly, different evolutionary scenarios
were proposed based on independent cladistic analyses, e.g.
Langer & Benton (2006), Ezcurra (2006), Sereno (2007b),
Irmis et al. (2007a), which attempted to sum up information
in order to understand better the interrelationships of early
dinosaurs.
(2) The dinosauromorph radiation
For most of the last century, it was accepted that dinosaurs
arose from ‘‘thecodont’’ precursors, either as a monophyletic
group or, more frequently (Fig. 2), in the form of inde-
pendent lineages (Huene, 1956; Colbert, 1964; Charig,

Fig. 1. Early images depicting some of the oldest putative dinosaurs. (A) Drawing of the slabs containing Saltopus elginensis, from
Huene (1910). (B) Skeletal reconstruction of Herrerasaurus ischigualastensis as mounted in 1965 for exhibition at the Universidad
Nacional de Tucumán, from Bonaparte (1997). (C) Skeletal reconstruction of Pisanosaurus mertii, from Bonaparte (1997). (D) Skeletal
reconstruction of Staurikosaurus pricei, from Colbert (1970). Scale bars: A = 5 cm; B-D = 10 cm.

Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
58
Fig. 2. Schemes of archosaur evolution depicting a polyphyletic Dinosauria. (A) Modified from Krebs (1974). (B) After Thulborn
(1975).
Attridge & Crompton, 1965; Romer, 1966). ‘‘Thecodonts’’,
as composed of non-crown-group archosaurs, and basal
members of both the bird and crocodile lines, are cur-
rently regarded as a paraphyletic group (Currie & Padian,
1997b; Benton, 2004). In his seminal paper on dinosaur
phylogeny, Gauthier (1986) applied the name Ornithosuchia
Huene, 1908, to designate a group composed of dinosaurs,
pterosaurs (including Scleromochlus), ornithosuchids, Euparke-
ria (questionably), and ‘‘Lagosuchus’’, a small archosaur from
the Middle Triassic of Argentina (Romer, 1971; Bonaparte,
1975; Sereno & Arcucci, 1994). That clade was supposed to
group all archosaurs that share a closer affinity to birds (within
Dinosauria) than to crocodiles, which were placed in its sis-
ter group Pseudosuchia (Parrish, 1997; Senter, 2005). More
recent work, however, excluded both Euparkeria (Benton &
Clark, 1988; Sereno, 1991a; Juul, 1994; Benton, 2004) and
ornithosuchids (Sereno, 1991a; Juul, 1994; Benton, 2004)
from Ornithosuchia, restricting the inclusivity, and perhaps
worthiness (Taylor, 2007) of the name. Indeed, alternative
names were later proposed for the bird line of Archosauria,
e.g. Avemetatarsalia Benton, 1999; Panaves Gauthier & De
Queiroz, 2001. The inclusivity of this group could be even
more reduced considering the labile position of pterosaurs,
sometimes regarded as basal archosaurs (Bennett, 1996) or
even outside Archosauria (Peters, 2000; Sobral & Langer,
2008). In this scenario, the non-dinosaur members of the
bird-lineage of Archosauria would only include Scleromochlus
taylori (a putative sister taxon to Pterosauria) from the Late
Triassic of Elgin (Sereno, 1991a; Benton, 1999) and the
so-called basal dinosauromorphs.
The name Dinosauromorpha was coined by Benton
(1985) to include dinosaurs, birds, and ornithosuchids, but
redefined by Sereno (1991a) to its current understanding,
which excludes ornithosuchids. The basal (non-dinosaurian)
Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
Max C. Langer and others
members of the group (Romer, 1971, 1972a, b; Arcucci,
1987) were for a long time known only from the Middle
Triassic Chañares Formation of Argentina (Rogers et al.,
2001). These small, gracile forms were grouped within
‘‘Pseudosuchia’’, but were soon recognized to have some
bearing on the origin of dinosaurs (Romer, 1972a, b), which
became evident with the works of Bonaparte (1975), Sereno
& Arcucci (1993, 1994), and Novas (1996). Those authors
identified typical dinosaur hind-limb traits on these taxa,
including a distally tapering fibula, an anterior ascending
process in the astragalus, a reduced calcaneum, a longer
metatarsus with reduced outer elements, and a straight
metatarsal V with reduced articulation area on the outer
surface of the lateral distal tarsal (see also Irmis et al.,
2007a; Brusatte et al., 2009). The taxonomy of the Chañares
dinosauromorphs has always been subject to some debate
(Bonaparte, 1975, 1995; Sereno & Arcucci, 1994; Arcucci,
1987, 1998, 2005), and five names entered the literature:
Lagerpeton chanarensis Romer, 1971; Lagosuchus talampayensis
Romer, 1971 (nomen dubium; Sereno & Arcucci, 1994);
Marasuchus lilloensis (Romer, 1972b, gen. Sereno & Arcucci,
1994); Lewisuchus admixtus Romer, 1972a (Arcucci, 1997); and
Pseudolagosuchus major Arcucci, 1987.
Recent discoveries (Fraser et al., 2002; Dzik, 2003; Irmis
et al., 2007a; Ferigolo & Langer, 2007) and interpretations
(Novas & Ezcurra, 2005; Ezcurra, 2006; Nesbitt et al., 2007)
suggest that basal dinosauromorphs were both more diverse
in terms of anatomy and inferred habits, and more widely
spread chronologically and geographically. Dromomeron romeri
and D. gregorii (Irmis et al., 2007a; Nesbitt et al., 2009) were
recognized in the Norian of western North America, which
also yielded Eucoelophysis baldwini. The latter taxon, first
described as a theropod dinosaur (Sullivan & Lucas, 1999),
was reassigned to a non-dinosaur dinosauriform position,
The origin and early evolution of dinosaurs
as either the sister taxon to Dinosauria (Ezcurra, 2006) or
forming a group with Silesaurus opolensis (Irmis et al., 2007a).
The latter form, collected in Carnian deposits of Poland
(Dzik, 2003; Dzik & Sulej, 2007), provided the greatest
breakthrough in the recent study of dinosaur origins. Its long
fore limbs suggest that the animal was at least facultatively
quadrupedal, while the edentulous front tip of its lower jaw
apparently bore a corneous beak. This atypical set of traits
revealed an unsuspected morphological diversity, hinting at
how incomplete was, and certainly still is, our knowledge of
the early stages of dinosauromorph evolution. In addition,
the record of Silesaurus opolensis extended the range of basal
dinosauriforms into the Late Triassic of Europe, a possibility
only hinted at before on the basis of controversial British taxa
such as Saltopus elginensis (Rauhut & Hungerbühler, 2000) and
Agnosphitys cromhallensis (Fraser et al., 2002). Further, since
the description of Silesaurus opolensis, newly and previously
described Norian forms have been considered closely related
to the taxon. This is the case for Sacisaurus agudoensis Ferigolo
& Langer, 2007, from the Caturrita Formation of South
Brazil, and a set of North American specimens (Nesbitt
et al., 2007), including material assigned to an unnamed
Silesaurus-like form from the Petrified Forest Member, Chinle
Formation, of New Mexico, and part of the original material
of Technosaurus smalli Chatterjee, 1984, from the Bull Canyon
Formation, Texas (Irmis et al., 2007b). The latter taxon
has been previously assigned to Ornithischia (Weishampel
& Witmer, 1990; Sereno, 1991b; Hunt & Lucas, 1994),
while Sacisaurus agudoensis might provide evidence that even
Silesaurus opolensis represents a basal member of that dinosaur
clade (Ferigolo & Langer, 2007).
The more complete non-dinosaurian dinosaurormorphs
form a series of outgroups to Dinosauria, and they give
clues about the origin of the clade (Ezcurra, 2006; Langer &
Benton, 2006; Yates, 2007a; Irmis et al., 2007a; Brusatte
et al., 2009). The long-held hypothesis of a more basal
position for Lagerpeton chanarensis (Novas, 1992b; Sereno
& Arcucci, 1993) was confirmed by independent studies
(Irmis et al., 2007a; Brusatte et al., 2009), which allocated
the genus Dromomeron as its sister taxon (Fig. 3A). Both
Lagerpeton and Dromomeron lack several apomorphic features
of Dinosauriformes such as a reduced medial lamina on the
pubis, an antitrochanter expanding into the ilium, a lesser
trochanter on the proximal femur, and a distal tibia bearing
a lateral groove and a squared distal articulation (Irmis et al.,
2007a; Brusatte et al., 2009). Within Dinosauriformes, most
studies (Novas, 1992b, 1996; Ezcurra, 2006; Irmis et al.,
2007a; Brusatte et al., 2009) place Marasuchus lilloensis as the
basalmost member of the clade (Fig. 3A). More derived forms
include Pseudolagosuchus major (Novas, 1992b, 1996) and its
possible senior synonym Lewisuchus admixtus (Arcucci, 1998,
2005). Along with the identification of further dinosauriforms
of equivalent grade (Dzik, 2003; Ezcurra, 2006), two
alternative phylogenetic scenarios were proposed (Fig. 3A).
Irmis et al. (2007a) suggested that Eucoelophysis and Silesaurus
form the sister clade to Dinosauria, which may also include
Pseudolagosuchus according to Nesbitt et al. (2007, p. 214).
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59
Ezcurra (2006), on the other hand, placed all these taxa in
a fully pectinated grade, where Pseudolagosuchus, Silesaurus,
and Eucoelophysis, are respectively closer to Dinosauria.
A somewhat intermediate view was adopted by Brusatte
et al. (2009), in which Pseudolagosuchus has a basal position,
and Lewisuchus forms, with other taxa, a more restricted sister
clade to dinosaurs (Fig. 3A). In any case, all or some of these
forms share with dinosaurs a number of apomorphic traits
absent in Marasuchus, e.g. longer pubic shaft; femur with
angular greater trochanter, ‘‘spike-like’’ lesser trochanter,
and prominent trochanteric shelf; distal tibia with laterally
expanded outer malleolus; astragalus with pyramid-shaped
anterior ascending process; and sigmoidal metatarsal IV
with deeper distal articular surface (Novas, 1996; Irmis et al.,
2007a; Brusatte et al., 2009).
Regardless of their status as a clade or ‘‘grade’’, these more
derived basal dinosauromorphs fill a gap (between Marasuchus
lilloensis and dinosaurs) in archosaur evolution. More
importantly, they fill that gap with the unsuspected diversity
of forms that have been informally called ‘‘silesaurids’’.
This group may just include Silesaurus, and forms such as
Sacisaurus and Technosaurus, which share with the Polish taxon
dental/jaw features possibly related to a more herbivorous
diet (Ferigolo & Langer, 2007; Irmis et al., 2007b), but it could
also encompass Lewisuchus, Pseudolagosuchus, and Eucoelophysis.
Although the basis for this assignment lies on shared traits
of the postcranium, there is no positive evidence that any
of these forms was a facultative/full quadruped as Silesaurus.
Yet, ‘‘herbivorous’’ teeth have been tentatively referred to
Eucoelophysis (Irmis et al., 2007a). The record of ‘‘silesaurids’’
and of the species of Dromomeron suggests that an extensive
radiation of non-dinosaurian dinosauromorphs preceded the
Late Triassic dinosaur diversification, and that parallel to the
first radiation of dinosaurs, that grade continued to flourish
after the Ladinian (Irmis et al., 2007a), extending their range
into the northern part of west Pangea (Fig. 3B).
II. PHYLOGENY AND SYSTEMATICS
The name Dinosauria was erected by Owen (1842) to
include three large terrestrial forms which he believed to
compose a distinct group of extinct reptiles (Torrens, 1992;
Padian, 1997a). In the following years, a sound concept
of Dinosauria was established by the proposition of several
classification schemes (Cope, 1866; Huxley, 1870; Marsh,
1882; Seeley, 1888). At that time, major taxa such as
Sauropoda and Theropoda (Marsh, 1878, 1881), as well as
Saurischia and Ornithischia (Seeley, 1888) were proposed.
These names gained acceptance in the 20th Century (Huene,
1932; Romer, 1956) and still represent the major dinosaur
subdivisions as currently understood (Fig. 4). However, for
most of the last century these different dinosaur groups, and
even some of their subgroups, were believed to have had
independent origins (Fig. 2) from ‘‘thecodont’’ precursors
(Huene, 1914, 1956; Colbert, 1964; Charig et al., 1965;
Romer, 1966; Reig, 1970; Krebs, 1974; Thulborn, 1975;
60 Max C. Langer and others

Fig. 3. Time-calibrated phylogenies and distribution of non-dinosaur Dinosauromorpha. (A) Recently proposed phylogenetic
hypotheses; dotted lines indicate ghost lineages; names applied as in Table 1. Position of Pseudolagosuchus in the phylogeny of Irmis
et al. (2007a) inferred from Nesbitt et al. (2007). (B) Geographic occurrence of taxa on a Late Triassic map redrawn from Blakey
(2006). Black silhouettes adapted from various sources.

Cruickshank, 1975, 1979). The monophyly of Dinosauria was
suggested by Bakker & Galton (1974) and Bonaparte (1975,
1976), firmly established by various pioneering cladistic
works (Paul, 1984; Gauthier & Padian, 1985; Cooper, 1985;
Brinkman & Sues, 1987), especially that of Gauthier (1986),
and represents a consensual hypothesis nowadays (Novas,
1989; 1996; Sereno et al., 1993; Sereno, 1999; Langer &
Benton, 2006; Irmis et al., 2007a).
(1) What makes a dinosaur?
Even if the monophyly of Dinosauria is consensually
accepted, the issue of which morphological traits characterize
the group continues to be debated (Novas, 1996; Langer &
Benton, 2006; Sereno, 2007b). Several putative dinosaur
apomorphies were proposed in a variety of studies dealing
with the phylogeny of the group, which frequently diverge
upon the distribution of these same characters. This is
epitomized by the continuing quarrel over one of the
diagnostic features mentioned by Owen (1842) in the original
proposition of the name: the number of vertebrae that
compose the dinosaur sacrum. In the following text, most
recent reviews of early dinosaur phylogeny (Novas, 1996;
Sereno, 1999, 2007a; Fraser et al., 2002; Benton, 2004;
Langer & Benton, 2006; Ezcurra, 2006; Yates, 2007a, b;
Irmis et al., 2007a; Brusatte et al., 2008a) are compared and
evaluated, in a search for the set of traits that typically
characterize the group. Obviously, a key point to set the
diagnosis of Dinosauria is to determine whether some of
the so-called basal dinosauromorphs actually belong to the

Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
The origin and early evolution of dinosaurs
Fig. 4. Generalized phylogeny depicting the position of Dinosauria and its main groups within Archosauria. Dotted lines indicate
major contentious placement of taxa; arrows indicate stem-based taxa; black circles indicate node-based taxa; names applied as in
Table 1; black silhouettes adapted from various sources.
group. As reviewed by Langer & Benton (2006, pp. 316-317),
various putative dinosaur apomorphies are seen in Silesaurus
opolensis. These might represent true dinosaur apomorphies
if the taxon is considered to represent a basal ornithischian
(Ferigolo & Langer, 2007). Yet, current orthodoxy points
towards the basal, non-dinosaurian position of Silesaurus, and
this hypothesis of relationships represents the template based
on which the unique dinosaur traits are discussed below.
Novas (1996) and Sereno (1999) respectively listed 17 and
18 characters as diagnostic for Dinosauria, while a modified
version of one of their characters (presence of three or more
sacral vertebrae) is the sole dinosaur apomorphy proposed
by Fraser et al. (2002). Langer & Benton (2006) critically
assessed these characters, questioning the apomorphic status
of several of them. Features related to the cranial anatomy
(Sereno & Novas, 1993) are particularly problematic because
most basal dinosaurs and, especially, basal dinosauromorphs
lack good skull material. Indeed, traits such as the lack of the
postfrontal bone, although typically absent in non-dinosaur
archosaurs and present in dinosaurs (see Irmis et al., 2007a,
char. 14), can not be considered an unambiguous dinosaur
apomorphy (Langer & Benton, 2006) given its equivocal
occurrence in most forms placed at the very origin of the
group. The same applies to other putative apomorphies of
the dinosaur skull, such as the dorsal overlap of the transverse
flange of the pterygoid by the ectopterygoid, and the lateral
exposure of the quadrate head (Langer & Benton, 2006); see
also Brusatte et al. (2008a, chars 10, 14, 38, 40, 67). The
status of other putative apomorphies of the dinosaur skull is
dependent on the position of Silesaurus opolensis, the cranial
material of which is reasonably complete (Dzik, 2003; Dzik
& Sulej, 2007). If not considered a dinosaur, some of its
Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
61
cranial traits, e.g. frontal participating in the supratemporal
fossa, are dismissed as dinosaur apomorphies. Yet, if its
less consensual position as a basal ornithischian is accepted,
these same traits continue potentially to represent dinosaur
synapomorphies. On the contrary, plesiomorphic traits in
the skull of Silesaurus such as a large post-temporal fenestra
supports its non-dinosaurian affinity, and helps to define a
reduced foramen-sized aperture (Fig. 5B) as apomorphic for
dinosaurs (Irmis et al., 2007a, char. 21). Other cranial features
(Langer & Benton, 2006, char. 12; Ezcurra, 2006, chars 4,
20; Yates, 2007a, chars 26, 29; Irmis et al., 2007a, chars 2,
25) suggested to represent possible dinosaur apomorphies,
pending the criteria used for character optimization, have an
erratic distribution among basal dinosaurs, and should not
be considered a priori diagnostic traits of the group. A likely
dinosaur apomorphy, related to the axial skeleton (Fig. 5C), is
the presence of epipophyses on the cranial cervical vertebrae
(Novas, 1996; Langer & Benton, 2006; Yates, 2007a; contra
Ezcurra, 2006). This feature was previously considered a
saurischian apomorphy, but more recently was recorded in
basal ornithischians (Novas, 1996; Langer & Benton, 2006;
Butler, Smith & Norman, 2007). Other putative apomorphies
of the dinosaur vertebral column listed by Yates (2007a, chars
129, 142) have an inconsistent distribution, and should not
be a priori considered as such.
As mentioned earlier, the increase in the number of
vertebrae that forms the dinosaur sacrum (from two to
more than two) continues to be listed as an apomorphy of
the group (Novas, 1996; Sereno, 1999; Fraser et al., 2002;
Ezcurra, 2006). Recently, as discussed by Langer & Benton
(2006), two main strategies of coding characters related to
this transformation have been employed; but see Novas
62 Max C. Langer and others

Fig. 5. The dinosaur Plateosaurus engelhardti. (A) Skeletal reconstruction (from Yates, 2003a), with indications of the better known
apomorphic traits of Dinosauria. (B) Occipital view of the skull (from Galton, 1985a) indicating (1) a foramen-sized post-temporal
fenestra. (C) Lateral view of a cervical vertebra, indicating (2) the presence of epipophyses. (D) Caudal view of the left humerus,
indicating (3) a long deltopectoral crest. (E) Lateral view of the left ilium, indicating (4) an open acetabulum and (5) an arched dorsal
margin. (F) Cranial view of the left femur, indicating (6) a femoral head inturned and distinctly offset from the shaft and (7) an
asymmetrical forth trochanter. (G) Proximal view of the left astragalus, indicating (8) an acute anteromedial corner, (9) a broader
ascending process, and (10) a reduced fibular articulation. (H) Cranial view of the distal tarsals, indicating (11) a proximally flat
lateral distal tarsal. All figured material refers to the mounted skeletons (GPIT I and III) of the ‘‘Sauriersaal’’ at Institut für Geologie
und Paläontologie, Tübingen (Weishampel & Westphal, 1986), except: B = SMNS 12949. Scale bars: A = 1 m; B-E, G-H = 5 cm;
F = 10 cm.

(1996) for a combined approach. Some (Fraser et al., 2002;
Rauhut, 2003; Ezcurra, 2006; Irmis et al., 2007a) adopted
a topographic criterion, simply considering the number of
sacral vertebrae, while others (Sereno et al., 1993, Sereno,
1999; Langer, 2004; Langer & Benton, 2006, Yates, 2007a)
attempted to recognize whether trunk or caudal elements
have been incorporated into the sacrum. Evidence for
a two-vertebrae sacrum within basal dinosaurs is limited,
and restricted to incomplete specimens (Langer & Benton,
2006; Yates, 2007a; Sereno, 2007b). On the contrary, the
sacrum of Silesaurus is clearly composed of three sacral
vertebrae (Dzik & Sulej, 2007). Accordingly, based on the
current evidence, and considering Silesaurus as closely related
but outside Dinosauria, the statement that dinosaurs are
apomorphic in having a sacrum composed of more than
two vertebrae is misleading. A more detailed approach that
attempts to recognize trunk or tail additions to the sacrum
may provide further information. In a few basal dinosaurs, i.e.
Saturnalia tupiniquim, Herrerasaurus ischigualastensis, Staurikosaurus
pricei, Guaibasaurus candelarienesis, and Eoraptor lunensis, the two
primordial sacral vertebrae are readily recognized based
on their much larger rib articulations. Other vertebrae
may be incorporated into the sacrum from either the
trunk (Herrerasaurus, Eoraptor) or the caudal (Staurikosaurus,
Saturnalia) series, but none has a conspicuous sacral rib,
compared to the primordial elements. Such a robust third
element is known in Silesaurus opolensis, and we agree with
Dzik & Sulej (2007) that it is borne by a trunk vertebra
added to the sacrum. Among the major dinosaur groups, all
theropods and ornithischians have trunk vertebrae added to
the sacrum, as is also the case in sauropodomorphs, except
for Plateosaurus (Yates, 2003c) and, possibly, Thecodontosaurus
(Yates, 2007a). Accordingly, even if a trunk vertebra added
to the sacrum is seen in most basal dinosaurs, the presence
of this character in Silesaurus dismisses its apomorphic status
for the group. On the other hand, the incorporation of
a caudal vertebra to the dinosaur sacrum seems more
restricted, absent in various basal forms (i.e. Herrerasaurus,
Eoraptor) and most basal sauropodomorphs (Yates, 2007a).
Indeed, the presence of caudosacral vertebrae is also not
accepted as a dinosaur apomorphy. It is evident that we
are dealing with a highly homoplastic character, possibly
affected by frame shift phenomena (Galton & Upchurch,
2000). It is also of misleading codification if one considers the
ambiguous condition of vertebrae that bore small transverse
processes/ribs that attach to the ilium and/or other sacral
transverse processes/ribs; compare Herrerasaurus in Novas
(1993) and Sereno (2007b). The increase in the number
of sacral vertebrae is, generally speaking, surely a typical
dinosaur trait. Yet, until more information, possibly derived

Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
The origin and early evolution of dinosaurs
from better preserved specimens of key taxa, is available,
the number of sacral vertebrae, and also the incorporation
of either trunk or caudal elements in the sacrum cannot be
unambiguously defined as dinosaur apomorphies. Besides,
Langer & Benton (2006) considered a dorsally expanded
cranial margin of the first primordial sacral rib as apomorphic
for dinosaurs. Similarly, this condition was also recognized in
Silesaurus (ZPAL Ab III/404/3), and can not be considered a
dinosaur apomorphy in the phylogenetic framework adopted
here.
Few characters of the pectoral girdle and limb have been
considered apomorphic for dinosaurs. This may indicate that
these parts of the dinosaur skeleton are not very modified
relative to the basic archosaur condition. Yet, it may also
reflect the lack of knowledge regarding these anatomical
elements, especially the forearm and hand, in the outgroups
to Dinosauria. This is particularly the case with the char-
acters related to the reduction of the outer digits of the
dinosaur manus (Gauthier & Padian, 1985; Novas, 1996;
Sereno, 1999). Indeed, dinosaur digit IV is always sube-
qual to or shorter than metatarsal III and never possesses
more than three phalanges, none of which is an ungual
(Langer & Benton, 2006). In addition, almost no dinosaur
is known to possess more than two phalanges in manual
digit V. On the contrary, manual digits IV and V of other
archosauromorphs are elongated elements with three or
more phalanges. More recently, Butler et al. (2007) claimed
that an enlarged grasping manus (with elongated pre-ungual
phalanges, prominent dorsal extensor pits and proximal
intercondylar processes), previously considered typical of Her-
rerasaurus ischigualastensis and theropods (Sereno et al., 1993;
Sereno, 1999), may also be apomorphic for dinosaurs, due
to its occurrence in basal ornithischians (Eocursor parvus and
heterodontosaurids). However, the manus is unknown in
non-dinosaur dinosauromorphs, and it is ambiguous at which
point of basal dinosauromorph evolution these modifications
occurred. Likewise, although no sternal plates have been rec-
ognized in basal dinosauromorphs, this may simply represent
a preservation bias (Padian, 1997b), and their occurrence as
paired ossifications (Sereno, 1999) can not be regarded as a
trustworthy dinosaur apomorphy.
In fact, the single feature of the pectoral skeleton accepted
by most previous studies as apomorphic for Dinosauria
appears to be a long deltopectoral crest (Fig. 5D), which
extends for more than 30-35% of the humeral length.
Besides, as noted by several authors (Yates, 2007a; Irmis
et al., 2007a; Brusatte et al., 2008a), contrasting with that of
pseudosuchians and Silesaurus opolensis, the deltopectoral crest
of dinosaurs is subrectangular, rather than subtriangular or
rounded. Yet, although lacking its proximal margin, the
deltopectoral crest of Marasuchus lilloensis (PVL 3871) seems
of the subrectangular type (Bonaparte, 1975), implying a
more inclusive distribution for that trait. Likewise, a shorter
forearm relative to the humerus can not be accepted a priori
as a dinosaur apomorphy (Irmis et al., 2007a), given that
a plesiomorphic longer forearm is retained in Herrerasaurus
ischigualastensis and Eoraptor lunensis (Langer et al., 2007b).
Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
63
Most novel traits of the early dinosaur skeleton are seen
in the pelvic girdle and limb. These were often related
to the acquisition of an improved bipedal gait (Bakker &
Galton, 1974), as typical of most basal members of the
group. Further, some authors, e.g. Bakker (1971) and Charig
(1972, 1984), have suggested that these traits represent key
features that allowed, or even promoted, dinosaur radiation
in Late Triassic times, while most other archosaurs were in
decline. Regardless of their evolutionary consequences (see
Sections IV.2,3), it is true that the dinosaur pelvic girdle and
limb bear various apomorphic traits. Indeed, about half of
the features presented by Novas (1996) and Sereno (1999)
as diagnostic for dinosaurs are related to those elements
(exclusive of the sacrum), and similar ratios are seen in other
recent works: four out of 11 in Langer & Benton (2006);
seven out of 11 in Ezcurra (2006); eigth out of 15 in Yates
(2007a); and 10 out of 14 in Irmis et al. (2007a). Obviously, the
fact that these anatomical parts are relatively well known in
basal dinosauromorphs facilitates the recognition of dinosaur
apomorphies.
Regarding the pelvic girdle, a perforated acetabulum
(Bakker & Galton, 1974; Novas, 1996; Ezcurra, 2006; Yates,
2007a), better described as a straight to concave ventral
acetabular margin of the ilium (Langer & Benton, 2006; Irmis
et al., 2007a; Brusatte et al., 2008a), stands in most recent revi-
sions as a valid synapomorphy of Saurischia plus Ornithischia
(Fig. 5E), but that is not the case of a brevis fossa/shelf in the
iliac postacetabular ala (Novas, 1996; Sereno, 1999; Fraser
et al., 2002; Benton, 2004; Yates, 2007a). Whereas a shelf
is also present in Marasuchus (Fraser et al., 2002; Langer &
Benton, 2006; but see Novas, 1996), a fossa is not only
seen in some basal dinosauriforms (e.g. Silesaurus), but is also
lacking in herrerasaurids (Novas, 1992b, 1993, 1996; Langer
& Benton, 2006). More recently, Ezcurra (2006) proposed
a straight to convex dorsal margin of the ilium (Fig. 5E)
as a dinosaur apomorphy. Indeed, basal dinosaurs lack a
dorsally excavated ilium, which seems to be typical of basal
dinosauromorphs (Sereno & Arcucci, 1993, 1994), although
not well preserved in some (e.g. Silesaurus; ZPAL AbIII/361).
On the contrary, other recently proposed apomorphies of
the dinosaur ilium are either highly homoplastic (‘‘long
preacetabular process’’; Yates, 2007a) or define a more
inclusive clade (‘‘acetabular antitrochanter present’’; Irmis
et al., 2007a), i.e. Dinosauriformes (see Sereno & Arcucci,
1994). Irmis et al. (2007a) also suggested that a transversely
compressed distal pubis is a dinosaur apomorphy, reversed
in sauropodomorphs, but the definition and distribution of
this feature is not so straightforward, as extensively discussed
by Langer & Benton (2006, p. 338).
Irmis et al. (2007a, char. 73) newly proposed that the pubic
process of the dinosaur ischium is apomorphic, because
‘‘separated from the ilial peduncle’’. In fact, the surface
connecting the iliac and pubic articulations of the ischium
is simply excavated in many basal dinosaurs, especially
theropods (Coelophysis rhodesiensis, QVM QG 1; Liliensternus
liliensterni, MB R 2175) and ornithischians (Scelidosaurus
harrisoni, BMNH1111; Scutellosaurus lawleri, UCMP 130580;
64
Butler et al., 2007). On the contrary, in forms such
as Marasuchus lilloensis (PVL 3870) and Silesaurus opolensis
(ZPAL AbIII 1228, 404/1) that excavation does not
reach the medial-most margin of the ischium, so that a
medially displaced sheet of bone remains, filling the space
between pubic process and iliac peduncle. This condition is
reminiscent of more basal archosaurs, in which the ischium
contributes significantly to the composition of the medial wall
of a non-perforated acetabulum. Herrerasaurus ischigualastensis
(PVL 2566) retains a much reduced medial sheet of bone, so
that the acetabular surface of the ischium can be considered
fully excavated, i.e. bearing the dinosaur apomorphy as
defined by Irmis et al. (2007a). On the contrary, the condition
among sauropodomorphs is variable (Yates, 2003c); e.g. in
Saturnalia tupiniquim (MCP 3846-PV), although an extensive
antitrochanter disrupts the clear observation of the character
(but see Liliensternus liliensterni, MB R 2175), the medial sheet of
bone occupies the space between that structure and the pubic
articulation. Accordingly, the status of the character defined
by Irmis et al. (2007a) awaits further investigation. Other
previously proposed apomorphies of the dinosaur ischium
include the presence of a reduced medioventral lamina
(Novas, 1996; Langer & Benton, 2006) and a proximal
dorsolateral sulcus (Yates, 2007a). Yet, both features are
clearly present in Silesaurus (ZPAL AbIII 361, 404/1), so
that their status as apomorphic for dinosaurs depends on the
contentious position of that taxon.
The femur is possibly the most scrutinized bone in the study
of early dinosaurs, with more than ten different characters
found as apomorphic for the group in the phylogenies revised
here. An inturned and subrectangular femoral head, that is
distinctly set from the shaft, has been considered among
the typical traits of dinosaurs by Bakker & Galton (1974)
and Gauthier (1986). Yet, this general state was poorly
dismembered into distinct and well-defined phylogenetic
characters, in order to evaluate the apomorphic condition of
each. Sereno (1999) defined an angular ‘‘greater trochanter’’
(i.e. nearly straight angle between the proximal articulation
and the long axis of the shaft) as a dinosaur apomorphy,
but that trait was also recognized in basal dinosauromorphs
(e.g. Pseudolagosuchus major, PULR 53; Ezcurra, 2006). This
structures a subrectangular femoral head, if the latter is
distinctly offset from the shaft, as diagnostic of dinosaurs
(Ezcurra, 2006, char. 231; Irmis et al., 2007a, char. 81;
Brusatte et al., 2008a, char. 132). That condition appears
along with an inturned femoral head (Fig. 5F), which can be
also considered a dinosaur apomorphy.
Irmis et al. (2007a) claim that the femoral head of dinosaurs
apomorphicaly bears a ligament sulcus and an asymmetrical
fossa articularis antitrochanterica, but these traits have also
been recorded in other basal dinosauromorphs (Novas, 1996;
Ezcurra, 2006). Likewise, the apomorphic condition of a
reduced medial tuberosity (Novas, 1996; Sereno, 1999)
and a prominent lesser trochanter (Novas, 1996) have
been dismissed by most recent studies (Langer & Benton,
2006; Ezcurra, 2006; Irmis et al., 2007a). Other features of
the femoral head were considered apomorphic reversals of
Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
Max C. Langer and others
Dinosauria (Ezcurra, 2006, char. 232; Irmis et al., 2007a,
char. 85; Brusatte et al., 2008a, char. 135), but depend
on character optimization. Besides, although reversed in
theropods, the presence of an asymmetrical fourth trochanter
(Fig. 5F) appears as a valid dinosaur apomorphy in most
recent reviews (Langer & Benton, 2006; Ezcurra, 2006;
Irmis et al., 2007a), and has been recently recorded also in
Guaibasaurus candelariensis (Bonaparte et al., 2007; contra Langer
& Benton, 2006) and Chindesaurus bryansmalli (GR 226; contra
Yates, 2007a).
Previously defined tibial traits such as the presence of a
cnemial crest (Novas, 1996; Sereno, 1999) and a transversely
expanded distal articulation (Novas, 1996; Benton, 2004)
are no longer believed to represent dinosaur apomorphies,
given their erratic distribution among basal dinosaurs and
dinosauromorphs (Langer & Benton, 2006; Ezcurra, 2006;
Irmis et al., 2007a; Brusatte et al., 2008a). Similarly, because
also seen in Silesaurus, a descending process of the tibia that
caudally overlaps the ascending process of the astragalus
is also not regarded apomorphic for dinosaurs. According
to Yates (2007a), a sub-quadratic distal tibia and a thinner
fibula may represent dinosaur apomorphies, because the
reverse condition is seen in Silesaurus. Yet, the record of
the dinosaur condition in Marasuchus lilloensis jeopardizes
that assumption. Accordingly, no unambiguous apomorphy
is currently referred to the dinosaur pelvic epipodium. In
addition, Ezcurra (2006) considered, under DELTRAN
optimization, a tibia subequal to the femur as apomorphic for
dinosaurs. Although the contrary was described for Silesaurus
opolensis (Dzik, 2003), a longer tibia is not only typical of
basal dinosauromorphs (Sereno & Arcucci, 1993; 1994;
Pesudolagosuchus major, PVL 4629), but was also retained in
basal ornithischians (Santa Luca, 1980; Butler et al., 2007).
Indeed, among basal dinosaurs, only saurischians consistently
bear a subequal or longer femur; but see Staurikosaurus pricei
(Colbert, 1970).
The tarsal joint has also been the source of several
anatomical traits believed to characterize dinosaurs. Yet,
this is not the case of an astragalar ascending process and a
lateral articulation between the calcaneum and the astragalar
anterolateral process (Sereno, 1999), which were recently
identified in other basal Dinosauromorpha (Novas, 1996;
Langer & Benton, 2006; Brusatte et al., 2008a). Yet, a reduced
fibular articulation (Langer & Benton, 2006; Brusatte et al.,
2008a), a broader ascending process (Yates, 2007a, char.
314), and an acute anteromedial corner (Irmis et al., 2007a)
apparently stand as apomorphies of the dinosaur astragalus
(Fig. 5G). On the contrary, some putative apomorphies
of the dinosaur calcaneum, such as a concave fibular
articulation (Novas, 1996) and a rudimentary medial process
(Sereno, 1999) have an erratic distribution among basal
dinosauromorphs, and can not be unambiguously considered
as such (Langer & Benton, 2006; Ezcurra, 2006; Irmis et al.,
2007a; Brusatte et al., 2008a). On the other hand, as far
as the condition in the outgroups to Dinosauria can be
accessed, a proximally flat lateral distal tarsal (Novas, 1996;
Langer & Benton, 2006; Brusatte et al., 2008a) stands as a
The origin and early evolution of dinosaurs
unique trait of the group (Fig. 5H). The single apomorphy of
the dinosaur metatarsus proposed in the discussed studies
of early dinosaur phylogeny is the so-called ‘‘sigmoid’’
metatarsal IV (Sereno, 1999), a condition given by the
lateral displacement of the distal part of the bone (Novas,
1996; Brusatte et al., 2008a). This condition is, however, also
seen in some basal dinosauromorphs (Novas, 1996; Ezcurra,
2006), and disregarded as a dinosaur apomorphy.
(2) Phylogenetic definitions: naming early dinosaurs
With the advent of Phylogenetic Nomenclature (De Queiroz
& Gauthier, 1990, 1992, 1994), systematists acquired an
unprecedented tool to define taxon names in explicit
phylogenetic context, setting their composition according
to given hypotheses. A drawback of this revolution was
the inflation of phylogenetic definitions for various names
(Benton, 2000), as readily recognized in a brief inspection
of Paul Sereno’s webpage TaxonSearch. Indeed, when dealing
with these names, authors currently have to state which
of the available definitions is adopted to translate them
into the phylogenetic nomenclature system. The priority
issue is expected to be settled with the publication of
the ‘‘companion volume’’ of the PhyloCode (Cantino & De
Queiroz, 2007). Yet, before this volume is published and,
more importantly, accepted by the scientific community as
the ‘‘Systema Naturae’’ of phylogenetic definitions, these
will no doubt continue to proliferate in an unordered way.
In the following paragraphs, the phylogenetic definitions
pertinent to the discussion of dinosaur origins are treated in
historical order and, in an attempt to emulate the ‘‘Principle
of Priority’’ (ICZN, 1999), those first proposed, with small
modifications added if absolutely required, are listed in
Table 1 and employed throughout the text.
Because Jacques Gauthier was involved in the study
of archosaurs, including dinosaurs, he presented some
phylogenetic definitions for related groups (Gauthier &
Padian, 1985; Gauthier, 1986) even before the publication of
the paper that set the theoretical foundation of Phylogenetic
Nomenclature (De Queiroz & Gauthier, 1990). Gauthier
& Padian (1985) provided a phylogenetic definition for
Ornithosuchia, while Gauthier (1986) explicitly defined
Saurischia and Theropoda. Problematic aspects of these
definitions include the use of supraspecific and/or informal
specifiers (e.g. birds, archosaurs, crocodiles, dinosaurs,
sauropodomorphs, Ornithischia) and their choice based on
the phylogenetic orthodoxy of the time. Instead, we believe
that, for the sake of precision, newly proposed phylogenetic
definitions should use minimal groups as specifiers, and
for historical coherence rely, as much as possible, on taxa
mentioned in the original definition of the names. In any case,
because first published, those definitions are adopted here
for the names in question (Table 1). Alternative phylogenetic
definitions for Saurischia (Padian & May, 1993; Padian,
1997d; Sereno, 1998; Holtz & Osmólska, 2004; Langer,
2004) just replace specifiers, either because these are more
specific (Padian, 1997d; Sereno, 1998) or are quoted in the
original proposition of the name (Langer, 2004). Yet, based
Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
65
on current phylogenetic hypotheses, these circumscribe the
same set of taxa as Saurischia sensu Gauthier (1986). Similarly,
alternative specifiers in later definitions of Theropoda are
more specific (Currie, 1997) and either more highly nested
(Sereno, 1998) or first named (Padian, Hutchinson & Holtz,
1999; Holtz & Osmólska, 2004). Again, their use does not
change the inclusivity of the group as defined by Gauthier
(1986).
Further phylogenetic definitions pertinent to the discussed
groups were proposed by Sereno (1991a), Novas (1992b),
and Padian & May (1993). Sereno (1991a) gave node-
based definitions for Ornithodira Gauthier, 1986, and
Dinosauromorpha Benton, 1985. These had to be slightly
modified (Table 1) to fit the logical basis of Phylogenetic
Nomenclature and the updated taxonomy of Sereno &
Arcucci (1994), but substitute definitions (Benton, 2004)
are redundant. Especially problematic are the stem-
based definitions of Dinosauromorpha (Sereno, 1991a,
2005; Benton, 2004) that use pterosaurs as the external
specifier, given the uncertain phylogenetic position of these
reptiles. In their current understanding, Ornithodira and
Dinosauromorpha differ only by the inclusion of Scleromochlus
taylori and possibly pterosaurs in the former. The least
inclusive Dinosauriformes was node-based defined when
first named by Novas (1992b). This was modified (Table 1)
to fit the taxonomy of Sereno & Arcucci (1994), but equally
requires no substitute definitions (Benton, 2004).
Apart from the equivocal list of taxa presented by Gauthier
(1986, p. 44; see Padian, 1997a), Novas (1992b) provided the
first phylogenetic definition of Dinosauria as ‘‘the common
ancestor of Herrerasauridae and Saurischia + Ornithischia,
and all of its descendants’’. This is in agreement with
the taxonomic orthodoxy of the time (Gauthier, 1986;
Brinkman & Sues, 1987; Benton, 1990; but see Gauthier
et al., 1989), according to which: (1) saurischians plus
ornithischians form a clade, contrary to the traditional view
that these arose independently from ‘‘thecodont’’ precursors;
(2) herrerasaurids, conventionally regarded as saurischian
dinosaurs (Reig, 1963; Colbert, 1970), are basal to that
clade. Indeed, in order to keep herrerasaurids as dinosaurs,
Novas (1992b) used the former as an internal specifier of
the latter. By contrast, Padian & May (1993) explicitly
restricted the use of Dinosauria to the clade composed of
Saurischia and Ornithischia, exclusive of ‘‘Herrerasaurus and
its allies’’. Despite the ‘‘priority’’ of Novas (1992b), the latter
concept gained almost unconditional acceptance since (e.g.
Sereno, 1998, 2005; Fraser et al., 2002) and is employed
here (Table 1). In any case, these alternate definitions
only circumscribe different groups if herrerasaurids are
placed outside the Saurischia + Ornithischia dichotomy,
a hypothesis not supported by most recent studies (see
below). Other authors (Holtz in Padian, 1997a; Olshevsky,
2000; Clarke, 2004) attempted phylogenetically to define
Dinosauria using taxa included in the original proposition
of the name. In this case, the best option may be using all
names mentioned by Owen (1842) in a node-based fashion,
and to define Dinosauria as ‘‘the most recent common
66 Max C. Langer and others

Table 1. Phylogenetic definition of names relevant in the context of early dinosaur evolution.

Name Phylogenetic definition

ORNITHODIRA ‘‘Pterosauria, Scleromochlus, Dinosauromorpha (including birds), and all descendants
Gauthier, 1986 of their most recent common ancestor’’ modified from Sereno (1991a),
node-based
DINOSAUROMORPHA ‘‘Lagerpeton chanarensis, Marasuchus lilloensis, Pseudolagosuchus major, Dinosauria (inc.
Benton, 1985 Aves), and all descendants of their most recent common ancestor’’ modified from
Sereno (1991a); node-based
DINOSAURIFORMES ‘‘The most recent common ancestor of Marasuchus lilloensis, Dinosauria, and all taxa
Novas, 1992b stemming from it’’ modified from Novas (1992b); node-based
SILESAURIDAE ‘‘All archosaurs closer to Silesaurus opolensis, than to Heterodontosaurus tucki and
new name Marasuchus lilloensis’’; stem-based
DINOSAURIA ‘‘All descendants of the most recent common ancestor of birds and Triceratops’’
Owen, 1842 Padian & May (1993); node-based
ORNITHISCHIA ‘‘Dinosaurs closer to Triceratops than to birds’’ Padian & May (1993); stem-based
Seeley, 1888
GENASAURIA ‘‘Thyreophora and Cerapoda and all descendants of their common ancestor’’
Sereno, 1986 Currie & Padian (1997a); node-based
NEORNITHISCHIA ‘‘All genasaurs closer to Triceratops than to Ankylosaurus’’ Sereno (1998); stem-based
Cooper, 1985
THYREOPHORA ‘‘All genasaurs closer to Ankylosaurus than to Triceratops’’ Sereno (1998); stem-based
Nopcsa, 1915
SAURISCHIA ‘‘Birds and all dinosaurs that are closer to birds than they are to Ornithischia’’
Seeley, 1888 Gauthier (1986); stem-based
HERRERASAURIA ‘‘All dinosaurs that share a more recent common ancestor with Herrerasaurus than
Galton, 1985b with Liliensternus and Plateosaurus’’ Langer (2004); stem-based
HERRERASAURIDAE ‘‘Herrerasaurus, Staurikosaurus, their most recent common ancestor, plus all its
Benedetto, 1973 descendants’’ modified from Novas (1992b); node-based
EUSAURISCHIA ‘‘The least inclusive group of Saurischia, containing Cetiosaurus and Neornithes’’
Padian et al. 1999 Langer (2004); node-based
SAUROPODOMORPHA ‘‘The clade including the most recent common ancestor of Prosauropoda and
Huene, 1932 Sauropoda and all of its descendants’’ Salgado et al. (1997); node-based
MASSOPODA ‘‘The most inclusive clade containing Saltasaurus loricatus but not Plateosaurus
Yates, 2007a engelhardti’’ Yates (2007a); stem-based
SAUROPODIFORMES ‘‘The least inclusive clade containing Mussaurus patagonicus Bonaparte & Vince,
Sereno, 2005 1979, and Saltasaurus loricatus Bonaparte & Powell, 1980’’ Sereno (2005);
node-based
SAUROPODA ‘‘The most recent common ancestor of Vulcanodon karibaensis and Eusauropoda and
Marsh, 1878 all of its descendants’’ Salgado et al. (1997); node-based
THEROPODA ‘‘Birds and all saurischians that are closer to birds than they are to
Marsh, 1881 sauropodomorphs’’ Gauthier (1986); stem-based
NEOTHEROPODA ‘‘Coelophysis, Neornithes, their most recent common ancestor and all descendants’’
Bakker, 1986 Sereno (1998); node-based
COELOPHYSOIDEA ‘‘All ceratosaurs closer to Coelophysis than to Carnotaurus’’ Sereno (1998);
Nopcsa, 1928 stem-based
AEROSTRA ‘‘Ceratosaurus nasicornis, Allosaurus fragilis and all the descendants of their most recent
Paul, 2002 common ancestor’’ modified from Ezcurra & Cuny (2007); node-based

ancestor of Megalosaurus, Iguanodon, and Hylaeosaurus, and all its
descendants’’. Again, according to the current phylogenetic
hypotheses, this definition circumscribes the same set of taxa
as that of Padian & May (1993).
Novas (1992b) also proposed a node-based definition for
Herrerasauridae, ‘‘emended’’ by Novas (1997a). Yet, both
definitions are incomplete and a modified version of them is
employed here (Table 1). There is no good reason to replace
that definition with a stem-based Herrerasauridae (Sereno,
1998; Benton, 2004), especially because this is equivalent to
Herrerasauria (see below). Further, Padian & May (1993)
provided a stem-based definition for Ornithischia, in a
fashion that matches its mutual exclusivity in relation to
Saurischia sensu Gauthier (1986). Subsequent definitions use
more specific (Sereno, 1998) and also more ‘‘traditional’’
(Weishampel, 2004; Norman, Witmer & Weishampel, 2004a)
specifiers, but are equally inclusive based on current
phylogenies. Although the use of taxa mentioned in the
proposal of Saurischia (e.g. Allosaurus, Camarasaurus) and
Ornithischia (e.g. Stegosaurus, Iguanodon) may have been more
desirable, all the previous definitions successfully translate
Seeley’s (1888) dichotomous understanding of Dinosauria

Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
The origin and early evolution of dinosaurs
into the Phylogenetic Nomenclature system. Likewise, it
could also be argued that the use of apomorphy-based
definitions for Saurischia and Ornithischia better represents
that original proposition, given that the groups were defined
on a character basis, i.e. opisthopubic and propubic pelves.
Yet, this is problematic because only the ornithischian pelvic
construction is apomorphic, whereas saurischians retain the
general morphology seen in more basal archosaurs.
Salgado, Coria & Calvo (1997) first proposed a
phylogenetic definition for Sauropodomorpha (Table 1).
Their node-based definition preceded that (stem-based)
given by Upchurch (1997b) by a couple of months, but
both suffer from using Sauropoda and Prosauropoda as
internal specifiers. Subsequent proposals attempt to replace
those taxa by more specific, and deeply nested specifiers in
either a node- (Sereno, 1998) or stem- (Galton & Upchurch,
2004; Sereno, 2007a) based fashion. Although lower rank
specifiers are desirable, the same level of precision can be
achieved using higher taxa that are, in turn, defined with
direct reference (or by typification) to those minimal groups.
Moreover, the adequacy of an either stem- or node-based
Sauropodomorpha (Upchurch, Barrett & Galton, 2007) is
minor in face of the primacy of the definition provided by
Salgado et al. (1997).
More recently, Langer (2004) defined a stem-based
Herrerasauria Galton, 1985b, and a node-based Eusaurischia
Padian, Hutchinson & Holtz, 1999. The former group is
potentially equivalent to Herrerasauridae sensu Sereno (1998),
but the node-based original definition of Herrerasauridae is
employed here. In that context, Herrerasauria (Table 1) can
allocate dinosaurs closely related to, but outside the clade
composed of Herrerasaurus plus Staurikosaurus. Eusaurischia,
on the other hand, was first proposed to designate the clade
composed of Sauropomorpha plus Theropoda (Padian et al.,
1999). This is as inclusive as the stem-based Saurischia under
certain phylogenetic schemes (Novas, 1996; Sereno, 1999),
but excludes basal forms such as Eoraptor and herrerasaurs
in alternative frameworks (Langer, 2004; Ezcurra, 2006)
and remains a potentially useful name (Table 1). Finally,
Silesauridae is here defined as a stem-based taxon that
includes all archosaurs closer to Silesaurus opolensis than to
Marasuchus lilloensis and Heterodontosaurus tucki. The latter
form was chosen to represent Dinosauria because of its
completeness (Santa Luca, 1980) and basal position within
Ornithischia (Butler et al., 2007), a group to which Silesaurus
has been tentatively related (Ferigolo & Langer, 2007).
III. DINOSAUR ‘‘TRAIL BLAZERS’’ IN SPACE,
TIME, AND EVOLUTIONARY CONTEXT
(1) The oldest dinosaurs and the rocks that contain
them
For most of the last century, except in a few important
cases (Huene, 1926; Colbert, 1989; Sereno & Novas, 1992;
Sereno et al., 1993), the knowledge of Triassic dinosaurs was
Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
67
based on incomplete and/or fragmentary skeletal remains.
In the last decade, however, various studies (e.g. Rauhut
& Hungerbühler, 2000; Langer, 2004; Parker et al., 2005;
Ezcurra, 2006; Nesbitt et al., 2007) revised those early records,
questioning the dinosaur affinity of several of them. On the
other hand, the discovery of a variety of more complete
basal dinosaurs (e.g. Langer et al., 1999; Bonaparte et al.,
1999, 2007; Yates & Kitching, 2003; Butler et al., 2007; Pol
& Powell, 2007a, b; Martinez & Alcober, 2009; Ezcurra,
2008), allowed a more reliable picture to emerge. As detailed
below, this accounts for the possible, but poorly supported
Middle Triassic origin of the group, its first radiation during
the Carnian, and the full establishment of the main dinosaur
groups from the Norian onwards.
Usually, the oldest dinosaurs (Galton, 2000; Langer,
2004) are considered as coming from the Ischigualastian
beds (Langer, 2005a) of northwestern Argentina and south
Brazil (Fig. 6). These respectively include the Ischigualasto
Sequence, Ischigualasto-Ischichuca depocenter, Bermejo
Basin (Stipanicic & Marsicano, 2002; Currie et al., 2009),
and the Santa Maria Supersequence, Paraná Basin (Zerfass
et al., 2003), the continental sedimentation of which filled
extensional rift basins related to the Gondwanides orogenesis
(Zerfass et al., 2004). Early works dated the Ischigualasto
and Santa Maria formations as Middle Triassic (Romer,
1960, 1962; Reig, 1961, 1963), but a Late Triassic age,
first proposed by Bonaparte (1966), has been supported by
most recent biostratigraphic studies (Ochev & Shishkin, 1989;
Lucas, 1998; Langer, 2005a, b). This was corroborated by the
radiometric dating of the ‘Herr Toba’ bentonite (Fig. 6C),
at the base of the Ischigualasto Formation (Rogers et al.,
1993), that provided a 40 Ar/39 Ar age of 227 ± 0.3 Mya. Yet,
following the discrepancy between U-Pb and 40 Ar/39 Ar dates
(Schoene et al., 2006) and other comparative parameters,
Furin et al. (2006) recalculated a date of 230.3-231.4 ± 0.3
Mya. This corresponds to the late Ladinian in most timescales
(Ross, Baud & Manning, 1994; Remane et al., 2000; Ogg,
2004; Ogg, Ogg & Gradstein, 2008), but recent works
(Muttoni et al., 2001, 2004; Gallet et al., 2003; Kent, Muttoni
& Brack, 2006; Kozur & Weems, 2007) assigned older ages
for the Carnian boundaries. In that context, and considering
the sedimentation rate of comparable rift basins (Rogers
et al., 1993; Currie et al., 2009), the dinosaur-rich sites of the
lower third of the Ischigualasto Formation can be placed in
the latest Carnian. Yet, the middle third of that stratigraphic
unit, that also yielded dinosaur remains, may rest within
the middle Norian. This was recently corroborated by the
dating of another bentonite, from above the middle sector
of the Ischigualasto Formation (Currie et al., 2009), which
provided a 40 Ar/39 Ar age of 217.0 ± 1.7 Ma (Shipman,
2004), recalculated as 219.4-220.4 ± 1.7 Mya (M. Ezcurra,
personal observations).
Ischigualastian dinosaurs (Fig. 6C) include Herrerasaurus
ischigualastensis, along with its possible synonyms Ischisaurus
cattoi and Frenguellisaurus ischigualastensis (Novas, 1993), Eoraptor
lunensis (Sereno et al., 1993), and Panphagia protos (Martinez
& Alcober, 2009), from the lower third of the Ischigualasto
68 Max C. Langer and others

Fig. 6. Tectonic and sedimentary settings of southwestern Pangea during the Middle and Late Triassic, with emphasis on the South
American dinosaur-bearing sequences (Zerfass et al., 2004; Veevers, 2005). (A) Idealized east-west cross section from Santa Maria
intraplate rift to the Cuyo back-arc rift and Gondwanides orogen. (B) Palaeogeographic reconstruction; note that the extensional
basins are perpendicular to the transtensional stresses. Abbreviations as follows: SLV, Sierra de la Ventana; CFB, Cape Fold Belt.
Gondwanides orogen in grey. (C) Stratigraphic charts of the Bermejo and Paraná Basins, depicting the dinosauromorph/putative
dinosaur record. Fm., Formation; HAZ, Hyperodapedon Acme Zone according to Langer et al. (2007c); Mys, million years before
recent. Asterisks indicate possibly coeval faunas in which the dicynodont Jachaleria occurs.

Formation, and Pisanosaurus mertii (Bonaparte, 1976) from the
middle third of that stratigraphic unit (Rogers et al., 1993),
as well as Staurikosaurus pricei (Colbert, 1970) and Saturnalia
tupiniquim (Langer et al., 1999) from the Hyperodapedon
Assemblage-Zone of the Santa Maria Formation (Langer
et al., 2007b). More recently, the discoveries of two new
herrerasaurids (Martinez & Alcober, 2007; Ezcurra & Novas,
2008), a Saturnalia-like animal (Ezcurra & Novas, 2008;
Ezcurra, 2008), and a probable basal theropod (Martinez,
Sereno & Alcober, 2008) have been announced from the
Ischigualasto Formation. Outside South America, dinosaurs
of similar age are much less conspicuous (Fig. 7). These
mainly include fragmentary remains from Gondwanan areas
such as the possible record of Saturnalia in the Pebbly Arkose
Formation (Cabora Bassa Basin), lower Zambezi Valley,
Zimbabwe (Raath, 1996; Langer et al., 1999), and part
of the specimens attributed to Alwalkeria maleriensis, from
the Lower Maleri Formation (Pranhita-Godavari Basin), in
central Peninsular India (Chatterjee, 1987; Remes & Rauhut,
2005). The record of dinosaurs in other coeval deposits
such as the Timesgadiouine Formation (Argana Basin),
in Morocco (Jalil, 1996, Gauffre, 1993), and the Isalo II
beds (Morondava Basin), in Madagascar (Flynn et al., 1999),
has been dismissed (Jalil & Knol, 2002; Flynn et al., 2008).
According to Langer (2005b) the Ischigualastian can be
traced into northern Pangea to encompass the Lossiemouth
Sandstone Formation, in northern Scotland. Yet, the only
putative dinosaur from those strata, Saltopus elginensis, has
doubtful affinities to the group (Rauhut & Hungerbühler,
2000; Langer, 2004).
All dinosaur osteological records from pre-Ischigualstian
strata have been questioned, including Spondylosoma abscon-
ditum (Galton, 2000; Langer et al., 2007c), from the Santa
Maria 1 sequence in south Brazil (Fig. 6C). Further occur-
rences of the group in strata of equivalent age, mainly
based on fragmentary European specimens (Huene, 1932),
have also been dismissed (Benton, 1986b; Norman, 1990;
Galton & Walker, 1996; Rauhut & Hungerbühler, 2000).
On the other hand, suggestions that dinosaurs were already
present in Middle Triassic times are backed up by two lines

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The origin and early evolution of dinosaurs 69

Fig. 7. Distribution of the main tetrapod-bearing deposits of Late Triassic age and their dinosaur record. (A) Chinle Formation and
Dockum Group, western USA; (B) Newark Supergroup, North American Atlantic coast; (C) Argana Basin, Morocco; (D) Jameson
Land, Greenland; (E) Fissure-filling and Rhaetian deposits, northwestern Europe; (F) Germanic Basin, Central Europe; (G) Khorat
Plateau, Thailand; (H) Bermejo Basin, Argentina; (I) El Tranquilo Group, Argentina; (J) Paraná Basin, Brazil; (K) Karoo basins,
south-central Africa; (L) Morondava Basin, Madagascar; (M) Pranhita-Godavari Basin, India. Late Triassic map redrawn from
Blakey (2006). Generalized black silhouettes (not at the same scale) adapted from various sources. Fm., Formation; Mb., Member;
Mbs, members.

of evidence: trackways and the stratigraphic calibration of
phylogenetic hypotheses. Indeed, if silesaurids are accepted
as an inclusive sister taxon to Dinosauria (Nesbitt et al., 2007,
p. 214; Brusatte et al., 2008a; contra Ezcurra, 2006), encom-
passing Middle Triassic forms such as Pseudolagosuchus and
Lewisuchus, then the dinosaur stem (although not necessarily
dinosaurs) minimally arose at the same time, i.e. the Ladinian
Stage. This is supported by evidence extrapolated from the
palaeoichnological record. Tracks suggest the presence of
dinosauromorphs in the Middle Triassic of France (Lockley &
Meyer, 2000), Italy (Avanzini, 2002), and Germany (Haubold
& Klein, 2002). Some German tracks may correspond to
dinosaurs, as is also the case for Middle Triassic footprints
from various stratigraphic units in Argentina (Melchor & De
Valais, 2006; Marsicano, Domnanovich & Mancuso, 2007),
including the Los Rastros Formation (Fig. 6C). Although
these may also represent basal dinosauriforms, the already
diversified and somewhat advanced fauna of saurischians
found in the superposed Ischigualasto Formation, provides
some basis to infer a Middle Triassic origin of dinosaurs.
In the scheme proposed by Langer (2005b), some tetrapod
assemblages of the Newark Supergroup (Olsen, Schlische
& Gore, 1989), in the North American Atlantic coast
(Fig. 7), albeit slightly younger than those of the Ischigualasto
and Santa Maria formations, may correspond to the Late
Ischigualastian. These include the faunas of the Wolfville
(Fundy Basin, Nova Scotia) and Pekin (Deep River Basin,
North Carolina) formations, the ornithischian records of
which (Galton, 1983b; Hunt & Lucas, 1994) were considered
unsubstantiated by Irmis et al. (2007b). In any case, during
post-Ischigualastian times, dinosaurs became more abundant
and widespread. Some of these taxa have been known for
over a century (Meyer, 1837; Cope, 1889), but the diver-
sity of Norian dinosaurs (Fig. 8) was greatly enhanced by

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70 Max C. Langer and others

Fig. 8. Skeletal reconstructions (from various sources), at approximately the same scale, of selected Carnian and Norian dinosaurs,
partially depicting the Late Triassic diversity of the group. Scale bar (lower left ) = 1 m.

last-decade discoveries, especially from South America and
South Africa, as outlined below.
Post-Ischigualastian dinosaur faunas in South America
include those of the Los Colorados, Laguna Colorada, and
Caturrita formations (Langer, 2005a). The latter strati-
graphic unit, in south Brazil (Fig. 6C), has yielded the
saurischian Guaibasaurus candelariensis (Bonaparte et al., 1999,
2007), as well as the ‘‘prosauropod’’ Unaysaurus tolentinoi
(Leal et al., 2004). ‘‘Prosauropods’’ are well known in the La
Esquina fauna of the Los Colorados Formation (Bonaparte,
1972). That stratigraphic unit covers the Ischigualasto For-
mation in northwestern Argentina (Fig. 6C), and includes
Riojasaurus incertus (Bonaparte & Plumares, 1995), Coloradis-
aurus brevis (Bonaparte, 1978), and Lessemsaurus sauropoides (Pol
& Powell, 2007a), along with theropods (Bonaparte, 1972)
such as Zupaysaurus rougieri (Arcucci & Coria, 2003; Ezcurra
& Novas, 2007a). In Patagonia, the Laguna Colorada For-
mation (El Tranquilo Group) has yielded the ‘‘prosauropod’’
Mussaurus patagonicus (Bonaparte & Vince, 1979; Pol & Powel,
2007b) as well as a heterodontosaurid ornithischian (Baez
& Marsicano, 2001). Other dinosaur-bearing gondwanan
deposits of similar age (Fig. 7) include the Lower Elliot For-
mation (Stormberg Group, Karoo Basin), in South Africa
(Knoll, 2005), and the Upper Maleri Formation, in penin-
sular India. The latter, along with the overlying Lower
Dharmaram Formation, has yielded a diversified, but still
undescribed fauna of basal saurischians (Kutty & Sengupta,
1989; Novas et al., 2006), which may include a Guaibasaurus-
like form (Kutty et al., 2007). Basal sauropodomorphs are
also well known in the Lower Elliot Formation, where
Melanorosaurus readi, Antetonitrus ingenipes, Blikanasaurus cromp-
toni, Eucnemesaurus fortis, Plateosauravus cullingworthi, and a yet
unnamed form (Yates, 2003a, 2007a, b, 2008; Yates &
Kitching, 2003) were recorded along with the ornithischian
Eocursor parvus (Butler et al., 2007) and possible theropod teeth
(Ray & Chinsamy, 2002).
In North Pangea, various Norian faunas of Europe and
North America yielded dinosaur records (Fig. 7). These
include the rich prosauropod fauna of the German Keuper,
where Efraasia minor occurs in the Middle Stubensand-
stein (Löwenstein Formation) of Baden-Württemberg, along
with Procompsognathus triassicus and other possible theropods
(Hungerbühler, 1998; Rauhut & Hungerbühler, 2000; Yates,
2003a). Specimens/species attributed to Plateosaurus are
much more widespread both geographically and stratigraph-
ically (Yates, 2003c; Moser, 2003; Weishampel et al., 2004),

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The origin and early evolution of dinosaurs
also occurring in the overlying Knollenmergel (Trossin-
gen Formation, and related stratigraphic units) of Baden-
Württemberg, Bavaria, Lower Saxony, and Saxony-Anhalt,
as well as in putative coeval faunas from France, Switzerland,
and Greenland (Jenkins et al., 1994; Galton & Upchurch,
2004). In addition, the Thuringian Knollenmergel has
yielded the ‘‘prosauropod’’ Ruehleia bedheimensis and the thero-
pod Liliensternus liliensterni (Rauhut & Hungerbühler, 2000;
Galton, 2001). The ‘‘lower’’ fissure-filling deposits of the
British Isles (southwest England and south Wales) are also
frequently regarded as Norian in age (Fraser, 1994; Ben-
ton & Spencer, 1995), although they might well spread
into the late Carnian and/or Rhaetian (Benton et al., 2000).
Among these, the Pant-y-ffynnon site, in south Wales, is
better known for its dinosaur fauna, which includes the
basal sauropodomorph Pantydraco caducus (Kermack, 1984;
Yates, 2003b; Galton, Yates & Kermack, 2007) and a small
theropod possibly related to Coelophysis/Syntarsus (Rauhut &
Hungerbühler, 2000). P. caducus was previously assigned to
the genus Thecodontosaurus, the type species of which (T.
antiquus) is also known from various other putatively coeval
fissure-filling deposits (Benton & Spencer, 1995; Benton
et al., 2000). In addition, the Cromhall Quary, in Avon, has
yielded the specimens assigned to Agnosphitys cromhallensis, the
dinosaur affinity of which is controversial (Fraser et al., 2002;
Langer, 2004; Yates, 2007a). Perhaps, the youngest dinosaur-
bearing deposits of the European Triassic are the Rhaetian
beds of Normandy (northern France), Somerset-Avon (south-
west England), Mid-Glamorgan (south Wales), and Belgium.
These include the indeterminate theropod ‘‘Zanclodon’’ cam-
brensis (Rauhut & Hungerbühler, 2000; Galton, 2005a), the
coelophysoid Lophostropheus airelensis (Ezcurra & Cuny, 2007),
sauropodomorphs like Camelotia borealis (Storrs, 1994; see also
Godefroit & Knoll, 2003), and the very unlikely record of
a stegosaur (Galton, 2005a; Irmis et al., 2007b). In addi-
tion, a possible theropod has been recovered recently from
the Rhaetian beds of Lipie Śla̧skie, Poland (Dzik, Sulej
& Niedźwiedzki, 2008). Isolated ‘‘dinosaur’’ teeth, mainly
assigned to Ornithischia, have also been reported exten-
sively from Norian-Rhaetian European strata (Weishampel
et al., 2004), none of which was recently confirmed (Butler,
Porro & Heckert, 2006; Irmis et al., 2007b). The ‘‘Eurasian’’
record of Norian-Raethian dinosaurs (Fig. 7) is completed by
the basal sauropodomorphs of the Nam Phong Formation,
Thailand, that include Isanosaurus attavipachi (Buffetaut et al.,
1995, 2000).
The record of Triassic dinosaurs in western USA was
recently reviewed by Nesbitt et al. (2007; see also Parker
et al., 2005; Ezcurra, 2006; Nesbitt & Chatterjee, 2008).
No compelling evidence of either sauropodomorphs or
ornithischians was found, and only coelophysoids were
positively identified, along with putative basal saurischians
(herrerasaurs) and basal theropods (Fig. 7). Given that
the Santa Rosa Formation ‘theropod’ (Heckert, Lucas &
Sullivan, 2000) was considered an indeterminate archosaur
(Nesbitt et al., 2007), the oldest dinosaur from western
USA, and possibly the oldest known neotheropod so far
Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
71
is ‘‘Camposaurus arizonensis’’, an indeterminate coelophysoid
from the Placerias Quarry (Bluewater Creek Member, base of
the Chinle Formation), northern Arizona (Hunt et al., 1998).
Younger records of coelophysoids include Coelophysis bauri
(Colbert, 1989; Colbert et al., 1992; ICZN, 1996; Spielmann
et al., 2007), the material described by Cope (1889) and
Padian (1986), as well as other specimens (Ezcurra, 2006;
Irmis et al., 2007a; Spielmann et al., 2007), along with some of
those attributed to Gojirasaurus quayi (Carpenter, 1997; Nesbitt
et al., 2007). All these come from Norian deposits referred
to the Chinle Formation (Petrified Forest Member and
‘‘siltstone member’’), in central New Mexico and Arizona,
and the Bull Canyon Formation (Dockum Group), in east
New Mexico and west Texas (Nesbitt et al., 2007). Among
non-theropod dinosaurs, whereas Caseosaurus crosbyensis (Hunt
et al., 1998) was regarded as an indeterminate dinosauriform
(Nesbitt et al., 2007), putative herrerasaurs occur in the
Petrified Forest Member (Chindesaurus bryansmalli) in Arizona,
as well as in the Bull Canyon Formation, which also yielded
a putative basal theropod (Nesbitt et al., 2007; Nesbitt &
Chatterjee, 2008). In terms of age, except for those of the
Placerias Quarry, all reliable dinosaur occurrences in the
Triassic of western North America are considered younger
than the Blue Mesa Member of the Chinle Formation, in
Arizona (Nesbitt et al., 2007), which has been radiometrically
dated as 219.2 ± 0.7 Myr (Irmis & Mundil, 2008).
In conclusion, although a Middle Triassic (Ladinian) origin
of dinosaurs might be hypothesized, the oldest definitive
records of the group date from about 230 million years ago.
This corresponds to the Carnian stage of the Late Triassic.
Radiometric dating of different levels of the Ischigualasto
Formation, Argentina (Rogers et al., 1993; Shipman, 2004)
suggests that after about 20 million years, i.e. within the latest
Triassic, a more diverse (Fig. 8), and specially more abundant
and widespread dinosaur fauna was already present (Benton,
1983a; Ezcurra & Novas, 2008), as represented by the Los
Colorados Formation and correlated assemblages from other
parts of the world (Fig. 7).
(2) The evolutionary tree of early dinosaurs
‘‘Early dinosaurs’’ are broadly understood here as all putative
representatives of the group collected from Ischigualastian
strata, as well as younger dinosaurs, the position of which
within Ornithischia, Theropoda, or Sauropodomorpha, is
yet to be firmly established (Table 2). These include reason-
ably well-known forms such as Herrerasaurus ischigualastensis,
Pisanosaurus mertii, Staurikosaurus pricei, Eoraptor lunensis, Saturna-
lia tupiniquim, and Panphagia protos, as well as more fragmentary
taxa (Huene, 1910, 1942; Chatterjee, 1987; Long & Murry,
1995; Bonaparte et al., 1999; Fraser et al., 2002; Langer,
2004; Nesbitt et al., 2007). Pisanosaurus has always been con-
sidered an ornithischian dinosaur (Thulborn, 1971; Galton,
1972; Bonaparte, 1976), while Herrerasaurus and Staurikosaurus
were assigned into the base of Saurischia by pre-cladistic
works (Reig, 1963; Benedetto, 1973; Galton, 1977), although
more specific affinities to either sauropodomorphs (Reig,
1970; Colbert, 1970; Van Heerden, 1978) or theropods
72 Max C. Langer and others

Table 2. Taxonomic assignment of ‘‘early dinosaurs’’, as recently given by different authors.

Taxon Proposed affinity

Agnosphitys cromhallensis Non-dinosaur; Fraser et al. (2002)
Dinosauria (partim); nomen dubium; Langer (2004)
Basal Theropoda; Yates (2007a)
Basal Sauropodomorpha (Guaibasauridae); Ezcurra (2008)
Aliwalia rex Eucnemesaurus fortis; Yates (2007a)
Alwalkeria maleriensis Basal Saurischia (partim); Remes & Rauhut (2005)
Caseosaurus crosbyensis Dinosauriformes Nesbitt et al. (2007)
Chindesaurus bryansmalli Herrerasauridae Irmis et al. (2007a)
Basal Theropoda; Yates (2007a)
Basal Saurischia (partim); Nesbitt et al. (2007)
Eoraptor lunensis Basal Theropoda; Sereno (1999); Ezcurra (2006)
Basal Saurischia; Langer (2004); Yates (2005)
Guaibasaurus candelariensis Basal Saurischia (Guaibasauridae); Bonaparte et al. (2007)
Basal Theropoda; Yates (2007a), Langer et al. (2007a)
Basal Sauropodomorpha; Ezcurra (2008)
Herrerasauridae Basal Theropoda; Novas (1996); Sereno (1999)
Non-dinosaur; Fraser et al. (2002)
Basal Saurischia, Langer (2004); Yates (2005); Ezcurra (2006)
Herrerasaurus ischigualastensis Herrerasauridae Novas (1992b)
Panphagia protos Sauropodomorpha Martinez & Alcober (2009)
Pisanosaurus mertii Ornithischia Sereno (1999); Butler et al. (2007)
Saltopus elginensis Dinosauriformes Rauhut & Hungerb ühler (2000); Langer (2004)
Saturnalia tupiniquim Stem-Sauropodomorpha; Langer & Benton (2006)
Basal Saurischia (Guaibasauridae); Bonaparte et al. (2007)
Sacisaurus agudoensis cf. Ornithischia; Ferigolo & Langer (2007)
Non-dinosaur; Brusatte et al. (2008a)
Silesaurus opolensis cf. Ornithischia; Ferigolo & Langer (2007)
Non-dinosaur; Langer & Benton (2006)
Spondylosoma absconditum Non-dinosaur; Galton (2000)
cf. Herrerasauridae; Langer (2004)
Staurikosaurus pricei Herrerasauridae Novas (1992b)
Teyuwasu barberenai Dinosauria (partim); nomen dubium; Langer (2004)

(Galton, 1973; Bakker & Galton, 1974) were also claimed.
On the contrary, early cladistic studies (Fig. 9) depicted
Staurikosaurus and Herrerasaurus basal to the Ornithischia+
Saurischia dichotomy (Gauthier, 1986; Brinkman & Sues,
1987; Benton, 1990; Novas, 1992b), thus outside Dinosauria
on its emerging monophyletic understanding (Gauthier
et al., 1989), whereas contemporaneous studies never ques-
tioned the ornithischian affinity of Pisanosaurus (Novas, 1989;
Sereno, 1991b). These investigations set the basis to future
research on basal dinosaur phylogeny, accepting the group
as a monophyletic entity solely composed of Ornithischia
and Saurischia, the latter including equally monophyletic
Sauropodomorpha and Theropoda. Besides, Novas (1992b)
placed Herrerasaurus and Staurikosaurus into a monophyletic
Herrerasauridae, a hypothesis almost never contested since.
During the early nineties, new discoveries from the
Ischigualasto Formation, including almost complete skele-
tons of Herrerasaurus ischigualastensis (Sereno & Novas, 1992,
1993; Novas, 1993, Sereno, 1993) and the first record of
Eoraptor lunensis (Sereno et al., 1993), were announced along
with a new hypothesis of basal dinosaur relationships. This
was advocated based on independent numerical analyses
performed by Sereno et al. (1993; see also Sereno, 1999)
and Novas (1996) that found nearly identical results (Fig. 9).
The Herrerasauridae was depicted as the sister-taxon of
Neotheropoda, while Eoraptor was considered the basal-most
theropod. Apomorphic traits supporting the theropod affin-
ity of Eoraptor and Herrerasauridae were given as including
caudally curved tooth crowns not expanded at the base, a
broad axial intercentrum, elongated humerus and manus,
deep extensor pits on the distal end of metacarpals I–III,
and narrow metacarpal IV, as well as by typical preda-
tory adaptations shared by herrerasaurids and theropods
(Fig. 10), e.g. intramandibular joint, craniomandibular joint
at about the same level as the tooth rows, and manual
digits II and III with elongated penultimate phalanges
and strongly curved unguals with enlarged flexor tubercles
(Langer & Benton, 2006; Sereno, 2007b; but see Butler et al.,
2007). More recently, Chindesaurus bryansmalli was described
as a herrerasaurid (Long & Murry, 1995; Novas, 1997a;
Sereno, 1999), a phylogenetic hypothesis accepted by most
authors up to the late nineties. However, the suggestions that
Chindesaurus forms a clade with either Herrerasaurus (Novas,
1997a) or Staurikosaurus (Sereno, 1999) were not supported by
recent studies (Langer, 2004; Nesbitt et al. 2007; Bittencourt

Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
The origin and early evolution of dinosaurs 73

Fig. 9. Main alternative phylogenetic hypotheses depicting the interrelationships of ‘‘early dinosaurs’’, modified from the cited
sources. Arrows indicate stem-based taxa and black circles node-based taxa. Names applied as in Table 1, not as in the referred
publications. Abbreviations as follows: O, Ornithischia; T, Theropoda; H, Herrerasauridae; E, Eusaurischia.

& Kellner, 2009), which place the North American taxon
basal to the more ‘‘typical’’ herrerasaurids.
Around the turning of the 20th Century, the description
of new basal dinosauriforms such as Saturnalia tupiniquim and
Agnosphitys cromhallensis, led to the proposal of novel phylo-
genetic hypotheses of basal dinosaur relationships (Fig. 9).
Langer et al. (1999) described Saturnalia as the basal-most
sauropodomorph, within alternative phylogenetic arrange-
ments depicting Herrerasaurus and Staurikosaurus as either
saurischians basal to the Theropoda+Sauropodomorpha
dichotomy, or as a monophyletic sister taxon to Dinosauria.
Fraser et al. (2002) described Agnosphitys as the sister taxon
to Dinosauria, favouring a position of Herrerasaurus outside
that clade, but these results were not replicated by any
quantitative analyses performed since then.
In a comprehensive analysis of basal theropod phylogeny,
Rauhut (2003) recovered Eoraptor lunensis and herrerasaurids
as basal theropods, as first proposed by Sereno & Novas
(1992). However, most subsequent studies, including some
focused on basal theropods (Yates, 2005; Smith et al., 2007),
contradicted that hypothesis of basal dinosaur relation-
ships. Yates (2003b) conducted a cladistic study of basal
sauropodomorphs, and found a new phylogenetic arrange-
ment among saurischians where herrerasaurids were con-
sidered the sister group of all other components of the
clade, termed Eusaurischia by Padian et al. (1999). Yates
(2003b) also found Saturnalia tupiniquim as the most basal
sauropodomorph, as previously claimed by Langer et al.
(1999) and mainly accepted since. New comprehensive anal-
yses by Langer (2004; see also Langer & Benton, 2006) inde-
pendently came to similar results (Fig. 9). These corroborated

Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
74 Max C. Langer and others

Fig. 10. Selected anatomical features of Herrerasaurus ischigualastensis, depicting a combination of apomorphic traits shared with
Neotheropoda (underlined) and plesiomorphic states relative to the Eusaurischia condition (non underlined). Asterisks indicate traits
also seen in basal ornithischians according to Butler et al. (2007). Skeletal reconstruction based on Sereno (1993). Scale bar = 10 cm.

the position of herrerasaurids as basal saurischians, adding
Eoraptor lunensis as the sister taxon to Eusaurischia, and
Guaibasaurus candelariensis as a basal theropod. Indeed, sev-
eral eusaurischian apomorphies are lacking in herrerasaurids
(Fig. 10) and/or Eoraptor, as exemplified by a short caudoven-
tral premaxillary process, a nasal that possesses a caudolateral
process and forms part of the dorsal border of the antor-
bital fossa, caudal cervical vertebrae longer than cranial
trunk vertebrae, a large medial-most distal carpal, a stout
metacarpal I with lateral distal condyle distally expanded,
a long metacarpal II relative to metacarpal III, and an
expanded distal end of the ischium (Langer & Benton, 2006).
Subsequent studies broadly agree with the above scenario
(Irmis et al., 2007a; Nesbitt & Chatterjee, 2008; Martinez
& Alcober, 2009), but differ in minor details (Fig. 9). In
a study focused on the non-dinosaurian affinity of the
putative coelophysoid Eucoelophysis baldwini (Sullivan & Lucas,
1999), Ezcurra (2006) placed herrerasaurids as non-theropod
saurischians, but Eoraptor as a basal theropod, sister taxon
of Neotheropoda. In a study of basal sauropodomorph phy-
logeny, Upchurch et al. (2007) placed both Herrerasauridae
and Eoraptor as non-theropod saurischians, but considered
the former group as the sister taxon of Eusaurischia. Yates
(2007a, b) expanded his previous studies, adding Agnosphitys
cromhallensis, Guaibasaurus candelariensis, Chindesaurus bryansmalli,
and Eoraptor lunensis to an analysis of basal sauropodomorphs.
The former three taxa were found as basal theropods, with
Guaibasaurus as the sister taxon of a clade including Chin-
desaurus plus Neotheropoda, and Agnosphitys as the most
basal theropod. Yet, the theropod affinity of Chindesaurus
was challenged by Irmis et al. (2007a), who supported its
more traditional relation to Herrerasaurus, both lying basal to
the sauropodomorph/theropod dichotomy, as also suggested
by Langer (2004) and Nesbitt et al. (2009). More recently, the
diversity of basal members of the sauropodomorph lineage
was increased by the discovery of Panphagia protos (Martinez
& Alcober, 2009) and the undescribed sister-taxon of Satur-
nalia tupiniquim (PVSJ 845; Ezcurra, 2008), both from the
Ischigualasto Formation, of Argentina. Further, Ezcurra
(2008) also included Agnosphitys and Guaibasaurus in that
dinosaur lineage. Indeed, Bonaparte et al. (2007) has already
proposed a close relation between Guaibasaurus and Saturnalia,
forming Guaibasauridae at the base of Saurischia. However,
Langer (2004) suggested the theropod, possibly coelophysoid,
affinity of Guaibasaurus (see also Upchurch et al., 2007; Langer,
Bittencourt & Schultz, 2007a; Bittencourt, 2008).
Several putative basal dinosaurs were never included in
numerical phylogenetic analyses, and their affinities are
open to scrutiny. Langer (2004) offered a comprehensive
summary of these records, but this has to be updated with
new information available since. Remes & Rauhut (2005)
reassessed the affinity of Alwalkeria maleriensis, first described as
a basal theropod (Chatterjee, 1987; Norman, 1990), but later
regarded as a dinosaur of uncertain (Novas, 1989, 1997a)
or eusaurischian (Langer, 2004) affinities. Those authors
found that the holotype represents a chimera, including
pseudosuchian and possible prolacertiform material, but also
saurischian specimens. Aliwalia rex Galton, 1985b, on the
other hand, previously regarded as a herrerasaurid (Galton,
1985b; Paul, 1988) or a dinosaur of dubious affinities (Sues,
1990; Galton & Van Heerden, 1998; Langer, 2004) was
shown to represent a junior synonym of Eucnemesaurus fortis,
therefore a basal sauropodomorph (Yates, 2007a). Nesbitt
et al. (2007) recently reviewed the status of the isolated
ilium previously assigned to Chindesaurus bryansmalli (Long
& Murry, 1995), which constitutes the holotype of Caseosaurus

Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
The origin and early evolution of dinosaurs
crosbyensis (Hunt et al., 1998) and considered the material
undiagnostic above Dinosauriformes. Other putative early
dinosaurs such as Saltopus elginensis, Spondylosoma absconditum,
and Teyuwasu barberenai, have not been studied recently.
Indeed, their uncertain affinities as proposed by Langer
(2004) are provisionally accepted here (Table 2).
In conclusion, recent cladistic analyses of basal dinosaur
relationships agree in various aspects, which are accepted by
most of the authors mentioned above: (1) dinosaurs represent
a monophyletic group exclusive of forms such as Lager-
peton chanarensis, Marasuchus lilloensis, Pseudolagosuchus major,
and Silesaurus opolensis; (2) Dinosauria is composed of two
main lineages, Saurischia and Ornithischia; (3) Pisanosaurus
mertii is a basal ornithischian; (4) Herrerasaurus ischigualastensis
and Staurikosaurus pricei belong into a monophyletic Her-
rerasauridae; (5) Eoraptor lunensis, Guaibasaurus candelariensis,
and herrerasaurids are saurischians; (6) Saurischia includes
two main groups, Theropoda and Sauropodomorpha; and
(7) Saturnalia tupiniquim and Panphagia protos are basal members
of the sauropodomorph lineage.
On the contrary, several aspects of basal dinosaur phy-
logeny remain controversial. These include the position
of herrerasaurids, Eoraptor lunensis, and Guaibasaurus cande-
lariensis as basal theropods or basal saurischians, and the
affinity and/or validity of various more fragmentary taxa
such as Agnosphitys cromhallensis, Alwalkeria maleriensis, Chinde-
saurus bryansmalli, Saltopus elginensis, Spondylosoma absconditum,
and Teyuwasu barberenai. Other equally incomplete forms
have been more thoughtfully studied, but while the affinities
of Aliwalia rex are better understood, Caseosaurus crosbyen-
sis continues to be problematic. In a reappraisal of the
methodologies employed in recent analyses of basal dinosaur
relationships, Sereno (2007b) highlighted that the lack of
consensus regarding the same phylogenetic problematics
is mainly due to differences in character/character-state
choice and codification among authors. Indeed, it seems that
more comprehensive studies, discussing these methodologi-
cal issues, are necessary to achieve a better understanding of
the phylogenetic relationships of basal dinosaurs. This is, in
turn, essential to recognize the patterns leading to their early
radiation and success during post-Triassic times.
(3) Geographical distribution of basal dinosaurs
The earliest records of dinosauromorphs and dinosauriforms
based on body fossils, and also most of the trustworthy
records of the earliest dinosaurs come from southern South
America, especially Argentina (e.g. Bonaparte, 1975; Sereno
& Novas, 1992; Sereno & Arcucci, 1993, 1994; Novas,
1992b, 1996; Langer et al., 1999; Galton, 2000; Rogers et al.,
2001; Langer, 2004; Ferigolo & Langer, 2007; Martinez &
Alcober, 2009). However, relatively few sites representative
of terrestrial ecosystems of that time are known (Hammer,
Collinson & Ryan, 1990; Lucas, 1998; Rogers et al., 2001;
Weishampel et al., 2004) and no biogeographic hypothesis
concerning the area of origin of the dinosaurian clades
can be robustly tested (Parker et al., 2005). Nonetheless,
whereas almost all south-Pangean tetrapod-bearing deposits
Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
75
of Carnian age (Langer, 2005b) bear undisputed, even if
inconspicuous dinosaur records, the north-Pangean scenario
is rather different, with no dinosaur positively identified in
coeval tetrapod assemblages. Accordingly, an admittedly
tentative scenario can be drawn, hinting at a southern
Pangean origin of dinosaurs.
Obviously, any biogeographical picture of dinosaur origins
has to be backed up by the current phylogenetic hypothe-
ses depicting the relationships of the basal members of
the group and its sister taxa. Accordingly, recent finds of
basal dinosauromorphs in Europe (Fraser et al., 2002; Dzik,
2003) and North America (Ezcurra, 2006; Irmis et al., 2007a;
Nesbitt & Chatterjee, 2008) came with new phylogenetic
proposals, and indicate that those animals had a broader
geographical and chronostratigraphic distribution than pre-
viously thought. Hypotheses that support an inclusive clade
of basal dinosauriformes (i.e. Silesauridae) as the sister taxon
to Dinosauria (Nesbitt et al., 2007; Brusatte et al., 2008a)
face the problem of a Ladinian ghost-lineage of ‘‘stem-
dinosaurs’’ (Fig. 3A), but are roughly in agreement with
the southern origin scenario. Spondylosoma absconditum, from
south Brazil, could fill that temporal gap, but its atypical
morphology and uncertain affinity (Langer, 2004) prevent
further scrutiny. The alternative pectinate topology (Ezcurra,
2006) overcomes the ghost-lineage problem, but suggests that
north Pangean taxa represent the immediate outgroups to
Dinosauria (Fig. 3A), jeopardizing the ‘‘out of south Pangea’’
model of dinosauromorph/dinosauriform/dinosaur radi-
ation. The record of Ladinian-Carnian ‘‘dinosauri-
form/dinosaur’’ footprints in various parts of the world (Mel-
chor & De Valais, 2006; Thulborn, 2006; Marsicano et al.,
2007) also hints at a broader distribution of these basal forms.
Late Triassic dinosaur records as a whole include body
fossils from Europe, North and South America, India,
Africa, and East Asia (Weishampel et al., 2004), as well
as putative tracks from Australia (Thulborn, 2000, 2006).
This is congruent with the geographic configuration of the
time (Fig. 7), when the Pangea Supercontinent and the lack
of extensive oceanic barriers would favour biotic expansion
(Shubin & Sues, 1991). Indeed, several non-dinosaur tetrapod
clades also achieved a widespread distribution during the
Late Triassic and Early Jurassic (Benton, 1993), but it
is important to note that no dinosaur clade had a truly
global distribution during Late Triassic times, especially
in the Carnian Stage (Nesbitt et al., 2007), even if only
the areas with tetrapod-bearing sites are considered. The
biogeographic patterns of early dinosaur radiation are, in
fact, better analyzed having the proposed subdivisions of the
group as a template.
The osteological record of Triassic ornithischians (Fig. 7)
is restricted to three Norian forms: the South African Eocursor
parvus (Butler et al., 2007), an unnamed heterodontosaurid
from Patagonia (Baez & Marsicano, 2001), and Pisanosaurus
mertii, from northwestern Argentina (Casamiquela, 1967;
Bonaparte, 1976), the latter of which may come from signifi-
cantly older deposits. Various other remains, mostly isolated
76
teeth, from either Europe (Godefroit & Cuny, 1997; Gode-
froit & Knoll, 2003) or North America (Chatterjee, 1984;
Hunt, 1989; Hunt & Lucas, 1994; Heckert, 2002, 2004)
had been assigned to the group. Along with footprints from
North America, Europe, and southern Africa, these may
hint at a broader Norian-Rhaetian geographical distribu-
tion of ornithischians. Yet, neither the isolated teeth nor
the footprints can be unequivocally assigned to the group
(Parker et al., 2005; Butler et al., 2006; Irmis et al., 2007b).
Accordingly, as discussed by Irmis et al. (2007b), ornithis-
chians do not seem to have been very diverse or abundant
through the Triassic, and certain hypotheses of relationship
(Sereno, 1991b, 1999; Xu et al., 2006) imply large gaps in
their fossil record. On the contrary, the usually accepted
basal position of Pisanosaurus is in accordance with its older
age, as is the possible basal position of heterodontosaurids
(Butler, Upchurch & Norman, 2008) and Eocursor (Butler
et al., 2007) in relation to other ornithischians. The suggested
heterodontosaurid affinity of Pisanosaurus (Bonaparte, 1976;
Galton, 1986; Crompton & Attridge, 1986; Butler et al., 2008)
implies a minimal ghost-lineage for Genasauria sensu Butler
et al. (2008), but is also in general agreement with a south
Pangean origin of ornithischians. Indeed, Laurasian occur-
rences of the group can not be confirmed before the Early
Jurassic (Irmis et al., 2007b), when basal thyreophorans occur
in North America, Europe, and Asia (Norman, Witmer &
Weishampel, 2004a; Irmis & Knoll, 2008). A different picture
emerges with the tentative placement of Sacisaurus agudoensis
and especially Silesaurus opolensis as the most basal ornithis-
chians (Ferigolo & Langer, 2007), but this hypothesis is still
to be backed up by numerical phylogenetic analyses. In any
case, the poorly documented early history of ornithischians
prevents any accurate biogeographic approach. According
to Irmis et al. (2007b), possible explanations for their rarity
in Late Triassic rocks (e.g. sample bias, differential environ-
mental occupation, systematic imprecision) are inconclusive.
Triassic saurischians have a much broader geographic
distribution (Rauhut & Hungerbühler, 2000; Langer, 2004;
Nesbitt et al., 2007). Indeed, basal members of the group,
and putative members of the theropod and sauropodomorph
lineages occur as body fossils in various Carnian beds
known from south Pangea (South America, southern Africa,
and India) as well as in Norian-Rhaetian deposits of
all continents except Australia and Antarctica (Fig. 7).
However, most records of ‘‘basal saurischians’’ are, in
fact, records of saurischians of uncertain affinities, and
only Eoraptor lunensis and herrerasaurs have been, under
certain phylogenetic hypotheses, positively placed basal to
Eusaurischia. Well-known herrerasaurids are restricted to
the South American Carnian (Langer, 2004; Bittencourt
& Kellner, 2009), including Herrerasaurus ischigualastensis and
Staurikosaurus pricei. The clade remains unidentified in the
relatively well-known post-Ischigualastian deposits of that
continent, hinting at its restricted stratigraphic distribution.
Yet, herrerasaurids have been identified in Norian beds
of western USA (Long & Murry, 1995; Hunt et al., 1998;
Irmis et al., 2007a; Nesbitt & Chatterjee, 2008), which would
Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
Max C. Langer and others
represent the younger records of the group. In addition,
given that the affinity of Chindesaurus bryansmalli to either
of the South American herrerasaurids has been questioned
(Langer, 2004; Bittencourt & Kellner, 2009), that North
American herrerasaur would most probably represent the
remnant of a lineage parallel to the typical members of the
group, and not a more derived outcome of that radiation.
On the contrary, the latter seems to be the case for the
specimen described by Nesbitt & Chatterjee (2008), which
bears herrerasaurid apomorphies.
If herrerasaurids and/or Eoraptor lunensis are treated as
theropods, the group would fit the ‘‘out of South Pangea’’
radiation pattern, with a well-known record of basal forms
in the Carnian of South America. In the alternative
arrangement (Langer, 2004), the oldest theropod, i.e. the
coelophysoid ‘‘Camposaurus arizonensis’’ (Nesbitt et al., 2007),
would not only come from North America, but also from
Norian-age deposits. If not filled by herrerasaurids and/or
Eoraptor this stratigraphic gap in theropod distribution is
unexpected, given the occurrence of basal members of
the sauropodomorph lineage in the Carnian (Langer et al.,
1999; Ezcurra, 2008; Martinez & Alcober, 2009), and
the abundance of both saurischian groups later in the
Triassic (Tykoski & Rowe, 2004; Galton & Upchurch, 2004).
Indeed, mainly represented by coelophysoids (but see Nesbitt
& Chatterjee, 2008), theropods become abundant during
Norian-Rhaetian times (Fig. 7), with body fossils recorded
in North America (Jenkins et al., 1994; Nesbitt et al., 2007),
Europe (Rauhut & Hungerbühler, 2000; Ezcurra & Cuny,
2007), Argentina (Arcucci & Coria, 2003; Ezcurra & Novas,
2007a), India (F. E. Novas, personal observations), and
perhaps South Africa (Ray & Chimsamy, 2002). The possible
theropod affinity (Yates, 2007a, b) of controversial Norian
taxa from Europe (Agnosphitys cromhallensis), North America
(Chindesaurus bryansmalli), Brazil (Guaibasaurus candelariensis),
and India (aff. Guaibasaurus) does not significantly change this
distribution pattern. In comparison to the more abundant
sauropodomorphs, osteological records of theropods are
lacking in Norian-Rhaetian deposits from southwest Asia
(Nam Phong Formation), suggesting that Pangean far-east
was first reached by the herbivorous/omnivorous branch of
Saurischia. Yet, this fossil assemblage is imperfectly known
(Buffetaut et al., 2000), and the absence of theropods might
simply represent a circumstantial sample bias. The scarcity of
theropod body fossils of Late Triassic age in southern Africa is
somewhat filled by ichnological evidence (Ellenberger, 1974;
Olsen & Galton, 1984; Raath et al., 1990), and also inferred
from their well-known Early Jurassic record (Raath, 1969;
Bristowe & Raath, 2004; Yates, 2005). Possible theropod
footprints are also known form Norian-Rhaetian deposits of
other parts of the world (Gatesy et al., 1999; Haubold & Klein,
2002; Demathieu & Demathieu, 2004; Thulborn, 2006), but
several of them have been questioned (King & Benton,
1996; Marsicano et al., 2007; Lucas, 2007; Nesbitt et al.,
2007), given their possible assignment to non-dinosaurian
dinosauromorphs. In any case, the overall record leads to a
scenario of low abundance of Carnian theropods, followed
The origin and early evolution of dinosaurs
by a significant Norian radiation, when the group occupied
most parts of Pangea.
Sauropodomorphs are surely the most abundant dinosaur
group of Triassic times. Basal members of the lineage, i.e.
Saturnalia tupiniquim and its allies, come from Ischigualastian
beds of South America (Langer et al., 1999; Da Rosa et al.,
2006; Ezcurra, 2008; Martinez & Alcober, 2009), and pos-
sibly southern Africa (Raath, 1996). This record could be
enhanced by the ‘‘prosauropods’’ of the Lower Maleri For-
mation referred to by Kutty & Sengupta (1989), but these
have not been mentioned in more recent studies (Kutty et al.,
2007), which refer the Upper Maleri ‘‘prosauropods’’ to aff.
Guaibasaurus. Assuming the above identifications as correct,
a southern radiation of ‘‘Saturnalia-like’’ forms may have pre-
ceded the Norian diversification of true sauropodomorphs.
The basal-most members of that group, Pantydraco caducus
and Thecodontosaurus antiquus (Yates, 2007a, b), come from
fissure-filling deposits of England and Wales of alleged Car-
nian age, but this occurrence better fits the much broader
distribution of later sauropodomorphs (Fig. 7). Indeed, the
Norian-Rhaetian record of the group excludes only Antarc-
tica, Australia, and continental North America (Vickers-Rich
et al., 1999; Galton & Upchurch, 2004; Nesbitt et al., 2007).
The former two areas, however, lack well-sampled tetrapod
faunas of that age, and the absence of sauropodomorphs
could represent a sampling bias. Indeed, the group can be
said to have had a nearly global Norian distribution, but this
was not uniform through time and space. Most of the basal,
non-Plateosauria (sensu Yates, 2007a) taxa were recorded
in Europe (Yates, 2003b,c; Galton & Upchurch, 2004;
Galton, 2007), while more derived forms are widespread
(Yates, 2007a, b; Galton & Upchurch, 2004; Leal et al.,
2004; Pol & Powell, 2007a). In this context, the lack of
sauropodomorphs in the Norian of continental North Amer-
ica (Nesbitt et al., 2007), though not in Greenland (Jenkins
et al., 1994), is intriguing. Indeed, this seems to represent a
true biogeographic pattern, given the abundance of well-
sampled tetrapod-bearing deposits of that age in the region.
The occurrence of the latest basal dinosauromorphs (Irmis
et al., 2007a) and probable herrerasaurs (Nesbitt & Chatter-
jee, 2008) in those faunas may also result from the causes that
drove this pattern. As for basal sauropods, previous studies
suggested that their distribution was initially restricted to
southeastern Asia, expanding throughout Pangea by the
Late Triassic-Early Jurassic (Gillette, 2003). Yet, recent phy-
logenetic hypotheses (Yates, 2007a, b; Smith & Pol, 2007;
Upchurch et al., 2007) have positioned south Pangean forms
like Antetonitrus ingenipes, Blikanasaurus cromptoni, Lessemsaurus
sauropoides, and Melanorosaurus readi, close to, or at the base
of Sauropoda. This suggests a wider distribution of early
members of the group, a pattern that seems to fit bet-
ter the footprint record (Wilson, 2005). In any case, most
Triassic ichnological evidence of sauropodomorphs was con-
sidered poorly substantiated (Lockley et al., 1994; Rainforth,
2002, 2003).
Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
77
IV. ECOLOGY OF THE DINOSAUR RADIATION
(1) The Triassic scene
During the Triassic, following the Late Permian maximum
coalescence of Pangea, most continental areas remained
forming a single landmass (Scotese, 2002; Golonka, 2002;
Blakey, 2006). Towards the end of the period, major rift zones
started to develop, especially along the Atlantic margins
of North America and North Africa, accounting for the
separation between Laurasia and Gondwana (LeTourneau
& Olsen, 2003; Golonka, 2007). Besides, the climate
experienced a trend towards higher instability compared
to Paleozoic settings (Holser & Magaritz, 1987; Kent &
Muttoni, 2003). The Triassic palaeoclimate was reviewed
in various landmark publications (Tucker & Benton, 1982;
Hallam, 1985; Parrish, 1993; Crowley, 1994; Golonka &
Ford, 2000), which suggest a warm period, when polar ice
caps were absent (Frakes, Francis & Syktus, 1992). Further,
a latitudinal zonation seems to have been present, with an
arid equatorial/tropical belt, a seasonally humid temperate
zone, and mainly humid higher latitudes (Hallam, 1985; but
see Fraser, 2006). In the second half of the period, a highly
seasonal (monsoonal) humid climate prevailed over various
parts of the supercontinent (Parrish, 1993). Triassic biotas
reflect the transitional nature of the time interval (Anderson &
Anderson, 1993; Fraser, 2006), particularly when terrestrial
tetrapod faunas are considered. The period starts with the
impoverished remaining diversity of the end-Permian mass
extinction (Benton, 2003), ending up with an essentially
modern fauna, that includes the first representatives of the
chelonian, lepidosaur, crocodilian, avian (in the form of
dinosaurs), and mammal lineages.
Part of the Triassic tetrapod diversity was inherited
from the Permian, when dicynodonts and limnarchian
temnospondyls reached their climax (King, 1988; Milner,
1993). These groups experienced a later diversification within
the Triassic, along with the first radiation of lineages of latest
Permian origin, like procolophonoids (Spencer & Benton,
2000), ‘‘protorosaurs’’ (Dilkes, 1998), archosaurs (Gower &
Sennikov, 2000), and cynodonts (Botha, Abdala & Smith,
2007). These tetrapod groups diversified through the Early
Triassic, composing the core of the Middle Triassic pre-
dinosaur terrestrial palaeocomunities; the ‘‘Kannemeyeroid
epoch’’ of Ochev & Shishkin (1989). An example of such
faunas is known from the Chañares Formation, Argentina
(Bonaparte, 1982; Rogers et al., 2001) that is dominated
by herbivorous cynodonts (Massetognathus) and dicynodonts
(Dinodontosaurus), along with predatory cynodonts (Chiniquodon)
and archosaurs (proterochampsids and ‘‘rauisuchians’’).
Nearly coeval faunas were recorded in Brazil, Russia, and
Southern Africa, further including a variety of limnarchians,
procolophonids, and rhynchosaurs (Ochev & Shishkin, 1989;
Lucas, 1998; Abdala & Ribeiro, 2003). In addition, as
previously discussed, the Chañares fauna also includes the
highest diversity of basal dinosauromorphs (Romer, 1971,
1972a, b; Bonaparte, 1975; Sereno & Arcucci, 1993, 1994).
78
The oldest known dinosaurs are recorded in a particular
faunal context, in which rhynchosaurs, especially the
genus Hyperodapedon, became dominant primary consumers
of various terrestrial faunas worldwide (Romer, 1962;
Benton, 1983b). These were recorded in the Hyperodapedon
Assemblage-Zone of the Santa Maria Formation (Fig. 11A),
the lower part of the Ischigualasto Formation, the
Lossiemouth Sandstone Formation, and the Lower Maleri
Formation (Langer, 2005b). Apart from rhynchosaurs and
the first dinosaurs, these faunas collectively encompass
limnarchian temnospondyls (Marsicano, 1999; Sengupta,
2003), various archosaurs such as proterochampsids (Price,
1946; Sill, 1967), ‘‘rauisuchians’’ (Huene, 1942; Alcober,
2000), poposauroids (Alcober & Parrish, 1997), aetosaurs
(Heckert & Lucas, 2002), phytosaurs (Chatterjee, 1978),
crocodylomorphs (Bonaparte, 1982; Ezcurra, Lecuona &
Irmis, 2008), and ornithosuchids (Benton & Walker, 1985),
carnivorous (Martinez, May & Forster, 1996; Bonaparte
& Barberena, 2001; Abadala & Gianinni, 2002) and
herbivorous (Bonaparte, 1962; Chatterjee, 1982; Hopson,
1985) cynodonts; as well as dicynodonts (Cox, 1965).
A similar faunal content was recorded in putatively coeval
faunas that lack rhynchosaurs and dinosaurs such as that
of Krasiejów, Poland (Dzik & Sulej, 2007), and the base of
the Irohalene Member (Timesgadiouine Formation, Argana
Basin), Morocco (Jalil, 1996). In fact, apart from the
appearance of some archosaur groups, and the dominance of
rhynchosaurs, the oldest dinosaur-bearing terrestrial faunas
are not significantly different from those of Middle Triassic
age.
Dinosaurs remain inconspicuous in younger faunas of
Norian age, as seen at the base of the Los Colorados
Formation (Caselli, Marsicano & Arcucci, 2001) and the
Caturrita Formation (Langer et al., 2007b), in South America,
and in some possibly coeval North American fossil assem-
blages (Langer, 2005b), i.e. Sanfordian faunas of the Newark
Supergroup; Camp Springs Member, Dockum Group; and
Popo Agie Formation (Huber, Lucas & Hunt, 1993; Lucas,
1998). These faunas include metoposaurids, procolophonids,
sphenodontians, Hyperodapedon, dicynodonts, traversodontid
and mammal-like cynodonts, as well as various pseudo-
suchians (aetosaurs, phytosaurs, poposauroids, and possible
‘‘rauisuchids’’). Later Norian deposits include a greater num-
ber of dinosaur records, within a slightly dissimilar faunal
context; the ‘‘Prosauropod’’ Empire of Benton (1983a).
As recorded from the top of the Los Colorados Forma-
tion, the fauna of La Esquina (Bonaparte, 1982; Caselli
et al., 2001) includes some of the oldest turtles, along
with crocodyliforms, ‘‘remaining’’ pseudosuchian lineages
(aetosaurs, ornithosuchids, ‘‘rauisuchians’’), and mammal-
like cynodonts (Fig. 11B). Putatively coeval faunas of other
parts of the world, especially South Africa (Anderson, Ander-
son & Cruickshank, 1998), Europe (Benton, 1994a), and
North America (Long & Murry, 1995), further include
various temnospondyls and phytosaurs, inconspicuous dicyn-
odonts (Dzik et al., 2008), the latest traversodontids (Hopson,
1984), as well as the first mammals (Lucas & Luo, 1993).
Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
Max C. Langer and others
(2) Lucky break?
Palaeoecological aspects of the early radiation of dinosaurs
and its correlation to Late Triassic extinctions and
corresponding biotic/environmental changes have been
addressed by various classical and more recent studies
(Colbert, 1958; Benton, 1983a; Charig, 1984; Olsen et al.,
2002; Tanner, Lucas & Chapman, 2004; Brusatte et al.,
2008a, b). Focus has been given to two inferred mass
extinction events, at the Carnian-Norian and Triassic-
Jurassic boundaries, and two alternative scenarios for the
rise of the dinosaurs, the so-called ‘‘competitive’’ and
‘‘opportunistic’’ models. Studies from the mid-late 20th
Century postulated that the replacement of various tetrapod
groups, notably pseudosuchians and therapsids, by dinosaurs
was a long-term affair driven by competition during the
Late Triassic (Cox, 1967; Charig, 1980; Bonaparte, 1982).
Its outcome would have been the dominance of dinosaurs
over terrestrial ecosystems from Norian/Jurassic onwards,
thanks to their ‘‘superiority’’ relative to the outcompeted
contemporary tetrapods, pushed to extinction. Usually, the
improved locomotory capability of the fully erect, bipedal
early dinosaurs was considered the most notable advantage of
the group (Charig, 1972, 1984), but their inferred advanced
physiology has also been mentioned (Bakker, 1971). From
the 1980s onwards, Benton (1983a, 1984, 1991) advocated
an alternative model, based on which the Triassic radiation
of dinosaurs was faster, opportunistically occupying adaptive
zones emptied by the extinction of rhynchosaurs, therapsids
(dicynodonts and some cynodonts), and pseudosuchians
(phytosaurs, aetosaurs, rauisuchians). More recently, Brusatte
et al. (2008a) demonstrated that Norian pseudosuchians
occupyied more morphospace and showed similar rates
of character evolution compared to dinosauromorphs/
dinosaurs. Indeed, this dismisses the classical ‘‘competitive’’
model, based on which those archosaurs were gradually
replaced by dinosaurs. Yet, the scenario seems to be more
complex in terms of patterns and timing of biotic turnovers,
as discussed below.
Of the two proposed Late Triassic extinction events
(Benton, 1986a, 1997), the end-Triassic is much better
documented in the literature than the end-Carnian, which
is often contested as minor or non-existent (Olsen & Sues,
1986; Olsen, Shubin & Anders, 1987; Hallam, 1990; Fraser
& Sues, 1994; Hunt et al., 2002). Classical studies reveal
the final demise of conodonts and a severe reduction
in the diversity of sponges, scleractinian corals, molluscs
(ammonoids, gastropods, and bivalves), and brachiopods in
the sea (Hallam, 1981; Raup & Sepkoski, 1982; Sepkoski,
1982, 1990), along with extinctions of insects (Benton, 1989)
and tetrapods on land (Olsen & Sues, 1986; Benton, 1994b).
Causes proposed for the end-Triassic mass extinction range
from sea level change (Hallam, 1990), to the impact of
one or more extraterrestrial bolides (Olsen et al., 2002)
and the establishment of the Central Atlantic Magmatic
Province (Marzoli et al., 1999). The latter two might have
led to an increase in the levels of atmospheric CO2 , and so
‘‘greenhouse’’ warming (McElwain, Beerling & Woodward,
The origin and early evolution of dinosaurs 79

Fig. 11. Reconstruction of two dinosaur-bearing fossil assemblages of the South American Late Triassic. (A) Alemoa fauna (Santa
Maria Formation), Carnian of south Brazil, depicting from left to right the aetosaur Aetosauroides sp.; the rhynchosaur Hyperodapedon
mariensis; the stem-sauropodomorph Saturnalia tupiniquim (group on background); the cynodont Prozoostrodon brasiliensis (in front), and
the herrerasaurid Staurikosaurus pricei. (B) La Esquina fauna (Los Colorados Formation), Norian of northwestern Argentina, depicting
on the left (from back to front), a group of the sauropodomorph Riojasaurus incertus; the ‘‘rauisuchid’’ Fasolasuchus tenax, and the
cynodont Chaliminia musteloides; on the right (from back to front), the crocodyliform Hemiprotosuchus leali, and the basal theropod
Zupaysaurus rougieri subduing the ornithosuchid Riojasuchus tenuiceps. Drawings by Jorge Blanco.

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80 Max C. Langer and others

Fig. 12. Distribution of medium to large sized terrestrial amniotes along the Late Triassic and Early Jurassic and the rise of dinosaurs.
Timeline from Gallet et al. (2003). Distribution of carnivorous (grey columns) and herbivorous/omnivorous (black columns) tetrapods
modified from Benton (1994a), according to Dilkes (1998), Abdala & Giannini (2002), Abdala & Ribeiro (2003), Thulborn & Turner
(2003), Lucas & Tanner (2005), Langer et al. (2007c), and Brusatte et al. (2008a). 1, Saturnalia tupiniquim; 2, Herrerasaurus ischigualastensis;
3, Guaibasaurus candelariensis; 4, Eocursor parvus; 5, Plateosaurus engelhardti; 6, Liliensternus liliensterni; 7, Massospondylus carinatus; 8, Vulcanodon
karibaensis; 9, Heterodontosaurus tucki; 10, Dilophosaurus wetherilli; 11, Scelidosaurus harrisoni. Silhouettes (roughly at the same scale) adapted
from various sources. Mys, million years before recent; Rhaet., Rhaetian.

1999). Yet, Tanner et al. (2004; see also Bambach, Knoll &
Wang, 2004) compiled evidence to reject what they call ‘‘the
myth of a catastrophic extinction at the Triassic-Jurassic
boundary’’. Indeed, as already hinted by some (Benton,
1994b; Cuny, 1995; Ezcurra & Cuny, 2007), although various
well-known Triassic amniote groups have no Jurassic record,
some might have gone extinct before the Triassic upper
boundary (Fig. 12).
The pioneering studies of Benton (1983a, 1986a, 1989,
1994b), which first challenged the long-term ‘‘competitive’’
model of dinosaur radiation, also advocated that the main
turnover of terrestrial faunas occurred at the Carnian-
Norian, rather than at the Triassic-Jurassic boundary.
This would have been characterized by the extinction of
various tetrapod groups, connected to climatic/floral changes
(Tucker & Benton, 1982). Problems related to that model
include: (1) new data suggest the survival, at least until
the initial stages of the Norian, of taxa believed to have
gone extinct at the end of the Carnian (Fig. 12); (2) lack
of synchronicity between the climatic/ floral changes of
north and south Pangea. Indeed, dicynodonts occur in
Norian faunas of South (Langer et al., 2007c) and North
America (Long & Murry, 1995), in the latest Triassic of
Poland (Dzik et al., 2008), and perhaps in much younger
assemblages as well (Thulborn & Turner, 2003). In addition,
the Caturrita Formation, of south Brazil, has yielded the
latest (Norian) remains of proterochampsid archosaurs and
rhynchosaurs (Langer et al., 2007c), while lagerpetonids
and herrerasaurids were recorded in the Norian of USA
(Irmis et al., 2007a; Nesbitt & Chatterjee, 2008; Nesbitt
et al., 2009). The diversity of chiniquodontid cynodonts, on
the other hand, has been reduced to a single genus of
Anisian-Carnian distribution in South America and southern
Africa (Abdala & Giannini, 2002; Abdala & Smith, 2009).
Accordingly, its absence in Norian strata does not represent
the demise of a well-established lineage, as is also the
case of single-genus ‘‘families’’ such as Pisanosauridae and
Scleromochlidae (Benton, 1994b). On the other hand, it
is important to stress that rhynchosaurs and dicynodonts
get less common in Triassic faunas after the Carnian. This
abundance shift, rather than their extinction, could indeed
provide some evidence for a biological crisis at the Carnian-
Norian boundary. In the marine realm, classical studies
suggested that invertebrate extinctions were not conspicuous
at the end of the Carnian (Simms & Ruffell, 1990), only
few groups suffering a moderate loss of diversity (Schäfer
& Fois, 1987; Smith, 1988). Yet, more recent data on
the so-called Reingraben Turnover suggest that a major

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The origin and early evolution of dinosaurs
restructuring of marine ecosystems occurred at the Carnian-
Norian boundary (Hornung & Brandner, 2005; Stanley,
2006; Hornung, Krystyn & Brandner, 2007b).
Despite Cornet & Olsen’s (1990) statement against
significant floristic changes across the Carnian-Norian
boundary, this is assumed in a climate-driven fashion by the
biotic substitution model of Benton (1983a). Most authors
agree on the occurrence of a monsoonal humid phase
(Fig. 12), minimally affecting the northern Tethyan realm
during the mid-late Carnian (Simms, Ruffel & Johnson,
1994; Hornung et al., 2007a). Yet, although a humid phase
was also recorded in the Late Triassic of other parts of
the world, it seems to have lasted until later within the
period in those areas. Prochnow et al. (2006) suggest that
approximately during the deposition of the Petrified Forest
Member of the Chinle Formation, presently considered of
Norian age (Nesbitt et al., 2007), western North America
was experiencing a period of increasing humidity. Similarly,
the Santa Maria Supersequence, in south Brazil, shows
the progressive replacement of an ephemeral anastomosed
fluvial-lacustrine system by a perennial braided fluvial system
throughout the Carnian and Norian (Holz & Scherer, 2000;
Zerfass et al., 2003), whereas the Norian upper third of the
Ischigualasto Formation bears evidence of a humidity peak,
based on plant taphonomy and the occurrence of argilitic
palaeosols (Colombi & Parrish, 2008). Besides, Dicroidium-
floras occur through the entire sequence, as is also the case
in other parts of south Pangea (White, 1990; Anderson &
Anderson, 1993), while bennettitaleans and conifers only
dominate after the Norian (Fig. 12).
What seems to occur during the Norian is a rise in the
abundance of dinosaurs (Fig. 12). These are inconspicuous
in Carnian faunas, representing about 5% of tetrapod
fossils collected in the Hyperodapedon Assemblage-Zone of the
Santa Maria Formation (Azevedo, Schultz & Barberena,
1990; Langer et al., 2007c) and the lower third of the
Ischigualasto Formation (Bonaparte, 1982; Rogers et al.,
1993). Carnivorous dinosaurs were proportionally more
abundant, representing nearly 40% of all terrestrial meat-
eaters, and half of the medium- to large-sized predators
of the latter assemblage (Rogers et al., 1993). By contrast,
dinosaurs represent from 25 to 60% of the terrestrial
tetrapods of classical Norian faunas (Benton, 1983a), notably
in South Africa (Kitching & Raath, 1984), Argentina
(Bonaparte, 1982), and Europe (Benton, 1994a). This is better
documented for saurischians, especially with the notable
radiation of ‘‘prosauropods’’. However, dinosaurs are still
minor components of possibly older Norian faunas such as
that of the Caturrita Formation (Langer et al., 2007c), where
they represent about 15% of the known diversity. Although
the total number of dinosaurs registered in the Norian is at
least three times higher than in the Carnian, Ezcurra & Novas
(2008) emphasize that dinosaur diversity in the Ischigualasto
Formation surpasses that of most Norian stratigraphic units
(Fig. 7). This may imply that their radiation in the latter
stage resulted in a more abundant and disparate (Brusatte
et al., 2008b), but not necessarily more diverse, dinosaur
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81
fauna, leaving the greater total diversity of Norian dinosaurs
to be explained based on the existence of more tetrapod-
bearing sites of that age. Yet, various main dinosaur groups
appear to have originated and/or radiated during the Norian,
as is the case for heterodontosaurids, coelophysoids, and
‘‘prosauropods’’.
Moreover, as discussed by Novas (1997b; contra Benton,
1983a), the Late Triassic dinosaur rise did not occur in
an empty ecospace. Instead, dinosaurs radiated during the
Ischigualastian despite the high diversity and abundance
of other tetrapods. Especially in the case of carnivores,
it is difficult to deny a degree of overlap in the use of
food resources by sympatric species of similar size (Glen &
Dickman, 2008); whereas modern analogues suggest that
feeding niche overlap is less significant among herbivores
(Plumptre, 1996; Begon, Harper & Townsend, 1996, p. 778).
In this context, it is possible to envisage strictly herbivorous
Triassic dinosaurs, which were not many (see Section
IV.3), using plant resources not previously exploited in
full. Yet, counterevidence is given by the non-overlap of
the rhynchosaur bearing Norian assemblages with younger
faunas of that age (Fig. 12), in which dinosaurs are more
abundant (Langer, 2005a). In this context, the Norian
radiation of herbivorous dinosaurs could be linked to
ecological release, given the extinction of rhynchosaurs (see
also Brusatte et al., 2008b). The corollary, as suggested
by Novas (1997b), is that competitive pressure of non-
dinosaur herbivores may account for the low abundance
of herbivorous dinosaurs in the Carnian.
Whatever the physical causes of the end-Triassic event,
direct ‘‘extermination’’ is unlikely to have been the only
agent of the extinctions. Instead, the exacerbation of
biotic interactions (including competition) in a changing
environment probably also played a major role. In that
context, the already abundant and diverse dinosaurs may
have had the key adaptations to succeed and expand,
within the shifting environment that drove various other
tetrapods towards extinction. This is the second explanation
offered by Brusatte et al. (2008a) accounting for the extinction
of pseudosuchians, and not dinosaurs, at the Triassic-
Jurassic boundary. It implies a circumstantial ‘‘superiority’’
of dinosaurs, while historical burden enforces the reference to
a fully erect gait (Charig, 1972) and/or advanced physiology
(Bakker, 1971; Ward, 2006) within the set of dinosaur
‘‘advantages’’. This explanation is preferred here, given that
the first is based on ‘‘chance’’ (Brusatte et al., 2008a), which
just reflects cases ‘‘when our knowledge does not suffice
for prediction’’ (Popper, 1959). In fact, as acknowledged
by Popper (1959): we may not infer from the fact that an
event is chance-like that its elements are ‘due to chance’.
This is because ‘‘chance’’ and lack of knowledge produce
the same signatures, and can not be set apart in practice. In
our understanding, opportunistic and competitive scenarios
of dinosaurs rise are not mutually exclusive, and competition
may have played an important role in that radiation episode.
Not as a long-term affair, but triggered by the physical events
ultimately linked to the end-Triassic extinction.
82
Although the Early Jurassic total diversity of dinosaur
genera is not significantly higher than the Late Triassic
(Wang & Dodson, 2006), it is usually accepted that another
pulse of dinosaur radiation followed the Triassic-Jurassic
extinction (Olsen et al., 2002). This is better measured by the
origin of certain lineages and exploitation of new ecological
roles (Fig. 12). Among saurischians, large carnivores (Ezcurra
& Novas, 2007a) and large quadruped herbivores (Yates &
Kitching, 2003) were already known in the latest Triassic.
In fact, the record of Zupaysaurus (Ezcurra & Novas, 2007a),
and studies of Brusatte et al. (2008b), falsifies the hypothesis
of theropod size increase due to ecological release after
the Triassic-Jurassic boundary (Olsen et al., 2002; Lucas &
Tanner, 2005). Yet the Early Jurassic saw the radiation
of the Dilophosaurus-clade (Smith et al., 2007) and further
acquisition of typical graviportal traits among sauropods
(Barrett & Upchurch, 2007). The radiation of ornithischians
was more notable, with heterodontosaurids attaining their
diversity peak and the origin of both neornithischians and
thyreophorans (Butler et al., 2007). The latter represents
the debut of quadrupedal armoured forms, within a
morphospace previously unoccupied by dinosaurs.
In conclusion, the radiation of dinosaurs comprises at least
three landmark moments (Fig. 12), mainly characterized
by early diversification (Carnian); increase in diversity
and, especially, abundance (Norian); and occupation of
new niches (Early Jurassic). As previously mentioned, the
Carnian diversification did not occur in an empty ecospace,
but despite the abundance and diversity of contemporary
tetrapods. The Norian increase in dominance might be
connected to climatic/ floristic changes and to the extinction
of herbivorous forms such as rhynchosaurs, but the timing of
these events needs further investigation. The subtle Jurassic
diversification, on the other hand, seems to have occurred in
the aftermath of an extinction event (Brusatte et al., 2008a).
Indeed, this might be an example of opportunistic radiation
into released ecospace (Benton, 1983a; Olsen et al., 2002;
but see Brusatte et al., 2008b). Obviously, this does not fit
the notion that the ‘‘end-Triassic’’ tetrapod extinctions were
scattered over the end of the period (Tanner et al., 2004).
Indeed, the lack of various tetrapod groups in Rhaetian beds
and the less than expected dinosaur diversity increase in the
Early Jurassic seem to justify this latter scenario, but the
‘‘diversity loss’’ of dinosaurs and other tetrapods during the
Rhaetian is likely to be due to sampling bias (Ezcurra &
Cuny, 2007). In any case, post-Triassic tetrapod biodiversity
can not be understood as the outcome of a single event, but
seems modeled by long-term coexistence of different groups
during the Late Triassic. Punctual events of environmental
change may have enhanced interaction among lineages,
leading to the extinction of some terrestrial forms. This was
probably topped by a final historical contingency at the
Triassic-Jurassic boundary, when dinosaur circumstantial
‘‘superiority’’ set the frame for the next 135 million years of
archosaur evolution.
Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
Max C. Langer and others
(3) Of legs and teeth: insights on the palaeobiology
of early dinosaurs
Non-crown-group archosaurs were all quadrupedal and
carnivorans (Charig, 1972; Parrish, 1986; Brusatte et al.,
2008a). This general pattern was retained in basal members
of the crocodile-line (Bonaparte, 1984; Sereno, 1991a),
although recent discoveries of bipedal (Nesbitt, 2007) and
herbivorous/omnivorous (Parker et al., 2005) pseudosuchians
notably amplified the disparity of these archosaurs. The
basalmost dinosaurs, instead, were all bipedal, but it is not
clear if this was also the case for more basal dinosauromorphs,
and several instances of reversion to full or facultative
quadrupedalism are known within the group (Padian, 1997c).
The feeding habit of basal dinosaurs is even more difficult to
assess. Basal dinosauromorphs have a rather unspecialized
dentition, but typical carnivores and herbivores occur early
in dinosaur evolution (Barrett, 2000).
As seen in the previous section, bipedalism is often consid-
ered a key dinosaur feature that, along with a fully erect pos-
ture, favoured the radiation of the group during Late Triassic
times. Yet, supporters of a polyphyletic origin of dinosaurs
suggested that some quadrupedal lineages, particularly
sauropods (Charig et al., 1965), have never had bipedal ances-
tors, evolving directly from quadrupedal basal archosaurs
(Fig. 2). The general acceptance of dinosaur monophyly, and
the identification of its sister taxa within gracile, and presum-
ably bipedal Middle Triassic archosaurs (Gauthier, 1986) set-
tled the new orthodoxy of originally bipedal dinosaurs, which
was in turn challenged by more recent data. This includes the
discovery of a potentially quadrupedal basal dinosauromorph
(Dzik, 2003), and new ichnological (Haubold & Klein, 2002)
and biomechanical (Fechner, 2006) interpretations that hint
at higher degrees of quadrupedalism among dinosaur precur-
sors. It is consensual that herrerasaurs, basal theropods, and
basal ornithischians were fully bipedal (Carrano, 2000; But-
ler et al., 2007), but the condition among sauropodomorphs
is less clear (Cooper, 1981; Barrett & Upchurch, 2007). In
any case, only if all evidence in favour of basal dinosauro-
morph/dinosaur quadrupedalism is accepted, and optimized
on a favourable phylogenetic framework (Fig. 13) does a
quadrupedal/facultative bipedal origin of dinosaurs emerge
as parsimoniously as a fully bipedal origin. Otherwise, the lat-
ter hypothesis is always favoured if herrerasaurs are regarded
as basal saurischians (Yates, 2003b; Langer, 2004; Ezcurra,
2006; Irmis et al., 2007a). In fact, a fully quadrupedal dinosaur
origin is consistently ruled out, given that basal members of
the sauropodomorph lineage, even if capable of walking on
all fours, must have relied on bipedalism for higher speed
locomotion (Christian & Preuschoft, 1996; Upchurch, 1997a;
Langer, 2003). Actually, this may be the case also of Silesaurus
opolensis, the unusual slender fore limbs of which may not
have endured the same amount of stress as the hind limbs did
(see Fariña, 1995). Evidently, more detailed biomechanical
studies of basal dinosauromorphs are needed to recognize if
dinosaurs originated from facultative or fully bipedal ances-
tors. At the moment, this is hampered by the scarce material
The origin and early evolution of dinosaurs 83

Fig. 13. Single hypothesis of basal dinosaur/dinosauromorph relationships in which the recognition of Lagerpeton, Silesaurus, and
sauropodomorphs as quadrupedal/facultative bipedal (grey silhouettes) allows an equally parsimonious reconstruction of this gait,
relative to a fully bipedal gait (white silhouettes), as ancestral to Dinosauria. White rectangles represent acquisition of bipedalism on
either a quadrupedal/facultative bipedal (C = convergences) or fully bipedal (A = apomorphy) scenario of dinosaur origins. Grey
rectangles (R = reversions) represent acquisition of facultative bipedalism/quadrupedalism on a fully bipedal scenario of dinosaur
origins. Silhouettes adapted from various sources.

assigned to most of these forms, specially regarding their fore
limb anatomy.
Classical scenarios of dinosaur dietary evolution (Charig
et al., 1965) depict independent origins of the major groups
from carnivorous ‘‘thecodonts’’. In fact, also in the cladistic
paradigm, typical ‘‘carnivorous teeth’’, i.e. pointed, caudally
curved, labiolingually flattened, unexpanded at the base, and
with finely serrated/denticulate keels, are usually accepted
as plesiomorphic for dinosaurs (Gauthier, 1986; Langer &
Benton, 2006, but see Barrett, 2000). Indeed, although more
conical in likely piscivorous forms (Sill, 1967; Hungerbühler,
2000), teeth of basal archosaurs (Ewer, 1965; Gower, 2003)
and pseudosuchians (Walker, 1964; Gower, 1999; Nesbitt,
2003) mainly fit into that pattern. Triassic exceptions
are pseudosuchians that bear modified dentitions towards
omnivory/herbivory/scavenging (Walker, 1961; Parker et al.,
2005). Given that known basal dinosauromorph teeth are all
labiolingually flattened, there is a good case that the above
described pattern is indeed ancestral to the group as a whole.
Dinosaur dentitions are, however, more heterogeneous.
Until recently, the dentition of non-dinosaur dinosauro-
morphs was inferred from the rather fragmentary tooth-
bearing bones referred to Lewisuchus admixtus (Romer, 1972a)
and Marasuchus lilloensis (Bonaparte, 1975). The isolated par-
tial jaw of Lewisuchus may not belong to the taxon, given
that it was found disarticulated in a concretion with other
archosaur taxa and seems larger in relation to the holotype
skeleton (PULR 01). The maxilla of Marasuchus (PVL 3870)
has been consensually attributed to the taxon (Sereno &
Arcucci, 1994), but its teeth are not well preserved enough
for an accurate inference of its diet. Most of the crowns fit
into the plesiomorphic pattern described above, but more
caudal elements seem slightly distally expanded at the base,
making them somewhat ‘‘leaf-shaped’’ (Bonaparte, 1975).
Yet, the lack of further dental modifications (see Barrett,
2000) precludes the assignment of an alternative diet to
Marasuchus, which probably fed on a variety of small ani-
mals. Lately, however, putative basal dinosauromorphs with
herbivorous adaptations have been discovered (Dzik, 2003;
Ferigolo & Langer, 2007; Irmis et al., 2007a). Silesaurus opolen-
sis and Sacisaurus agudoensis bear an edentulous beak in the
lower jaw, plus dental features usually associated with a
more herbivorous diet in ‘‘prosauropods’’ and ornithischians
(Galton, 1984; Crompton & Attridge, 1986; Sereno, 1991b),
but lately given as evidence of omnivory (Barrett, 2000; Irmis
et al., 2007b). Accordingly, although a full herbivore ancestry
of dinosaurs can be dismissed, there is some evidence that a
strictly carnivorous origin was also not the case.
Barrett (2000) comprehensively reviewed early dinosaur
feeding habits, remaining unsure about the ancestral
diet of the group. There is full agreement, however,
on the carnivorous habits of theropods and herrerasaurs
(Barrett, 2000; Bittencourt, 2008), while basal ornithischians
(Irmis et al., 2007b) and typical ‘‘prosauropods’’ (Barrett
& Upchurch, 2007) were most probably omnivorous. The
condition in some small-sized basal saurichians such as
Eoraptor lunensis and Saturnalia tupiniquim is more uncertain.
Eoraptor bore ‘‘leaf-shaped’’ rostral teeth, but its diet was most
probably still based on small animals. On the other hand,
most teeth of Saturnalia are ‘‘leaf shaped’’, and this animal
is frequently referred to as bearing herbivorous adaptations
(Barrett & Upchurch, 2007). However, the retention of finely
serrated tooth keels (Yates, 2003b) suggests that Saturnalia
was more carnivorous than any basal sauropodomorph.
The optimization of dental patterns into current
hypotheses of early dinosaur phylogeny reveals various
alternative scenarios. When forms with less modified
dentition such as Marasuchus and Eoraptor are considered
carnivores, the reconstruction of the ancestral dinosaur
feeding habit is ambiguous in most cases. Yet, if Eoraptor
plus herrerasaurs are considered basal theropods (Sereno,
1999), and Silesaurus opolensis and Sacisaurus agudoensis placed
in the sister clade to Dinosauria, then omnivory can be

Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
84
Fig. 14. Hypotheses of basal dinosaur/dinosauromorph rela-
tionships in which either omnivory (A) or carnivory (B) is
unambiguously reconstructed as ancestral to Dinosauria. White
and black silhouettes/circles, respectively, indicate omnivorous
and carnivorous taxa/optimizations. Silhouettes adapted from
various sources.
unambiguously regarded as plesiomorphic for the group
(Fig. 14A). On the contrary, if Silesaurus and Sacisaurus are
considered basal ornithischians (Ferigolo & Langer, 2007),
and either Eoraptor or herrerasaurs basal saurichians (Langer
& Benton, 2006), then dinosaurs are plesiomorphically
carnivores (Fig. 14B). Yet, it is important to stress that the
plesiomorphic dinosaurs tooth morphology does not strictly
compare to that of the typically carnivorous herrerasaurs
and theropods, or to the omnivorous/herbivorous pattern of
ornithischians and ‘‘prosauropods’’. Teeth of basal forms as
Marasuchus, Saturnalia, and Eoraptor, are less specialized, and
this was most probably also the case for the dinosaur common
ancestor, whether it was a strict carnivore or able to include
plant material in its diet. In this context, each major dinosaur
group seems to have independently acquired its typical
dental traits, in some cases along with a significant increase
in size. Indeed, the omnivorous/herbivorous dentition of
basal ornithischians and sauropodomorphs are sufficiently
different to preclude a common origin (Barrett, 2000). On the
contrary, the carnivorous teeth of herrerasaurs and theropods
are primary homologous (Bittencourt, 2008), and can be used
as evidence for the nesting of the former within the latter
group. Alternatively, they might have arisen independently,
as adaptations to the predatory habits of these animals
(Langer & Benton, 2006).
The skin of basal dinosaurs has always been thought as
scaly, but this view was recently challenged by the discovery
Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
Max C. Langer and others
of Tianyulong confuciusi (Zheng et al., 2009), which provides
the second evidence of an ornithischian with integumen-
tary filaments (see also Mayr et al., 2002). Its placement
within Heterodontosauridae (Zheng et al., 2009), i.e. the sis-
ter clade of all other ornithischians except Pisanosaurus (see
Section V.1), shows that at least some very basal ornithis-
chians bore integumentary filaments. Accordingly, if these
are actually homologous to the ‘‘protofeathers’’ of theropod
dinosaurs, the most recent common ancestor of saurischi-
ans and ornithischians would also likely bear this kind of
epidermal coverage. Witmer (2009) is cautious about the
dermal or epidermal origin of the integumentary structures
of Tianyulong, and their homology with those of theropods.
Yet, further research demonstrating the epidermal origin of
those filaments, a hypothesis currently supported by their
hollow structure (Zheng et al., 2009), would suggest that early
dinosaurs also bore integumentary filaments. If it was suffi-
ciently abundant, the coverage could play a thermoregulatory
function (insulation), probably implying a higher thermal
inertia (Regal, 1975, 1985; Unwin, 1998; Wu et al., 2004).
V. OUTCOMES OF A RADIATION
(1) Early ornithischian evolution
Although the concept of Ornithischia was defined only some
twenty years later (Seeley, 1888), the first dinosaur classifi-
cation schemes already congregated most members of the
clade we now know within a ‘‘natural group’’, e.g. Orthopoda
Cope, 1866. Indeed, ornithischian monophyly remains one of
the few uncontroversial issues in dinosaur taxonomy (Nopcsa,
1923, 1928; Huene, 1956; Romer, 1956, 1966; Steel, 1969;
Thulborn, 1971, 1972; Galton, 1972; but see Maryańska,
1977), having been fully corroborated by cladistic studies
(Sereno, 1999; Butler, 2005; Butler et al., 2007, 2008). Tradi-
tional ornithischian diagnostic traits such as the opisthopubic
pelvis (Seeley, 1888) and the predentary bone (Marsh, 1894)
can not be unambiguously considered apomorphic for the
group, because their occurrence in the suggested basalmost
ornithischian, Pisanosaurus mertii, is equivocal (Butler et al.,
2008). Yet, various other features have been accepted as
diagnostic for the clade, most of which correspond to modi-
fications of the teeth and tooth-bearing bones, related to the
acquisition of a more herbivorous diet. These include the
presence of a buccal emargination on the maxilla, into which
cheek musculature may have attached, and several changes in
the shape and arrangement of the teeth, as recently reviewed
by Irmis et al. (2007b) and Butler et al. (2008).
Five main ornithischian groups are traditionally
recognized: Stegosauria, Ankylosauria, Ornithopoda, Pachy-
cephalosauria, and Ceratopsia (Thulborn, 1972), but the rela-
tionships among these lineages remained ambiguous until the
first cladistic analyses of the group were performed (Sereno,
1984, 1986; Norman, 1984; Maryańska & Osmólska, 1985;
Cooper, 1985). Among these, the view advocated by Sereno
(1986, 1991b, 1999) was highly influential for nearly two
The origin and early evolution of dinosaurs
decades, during which few other phylogenetic studies focus-
ing on the basal radiation of Ornithischia were performed.
Sereno (1998, 1999) recognizes two main ornithischian splits:
Thyreophora and Neornithischia (Table 1). The former
includes Stegosauria and Ankylosauria, whereas Ceratop-
sia and Pachycephalosauria compose Marginocephalia, the
sister clade to Ornithopoda within Neornithischia. This gen-
eral scheme is accepted by most recent studies (Xu et al.,
2006; Butler et al., 2007, 2008), but details of basal forms
nesting within each major group remain controversial.
Sereno (1986) combined neornithischians and thyreo-
phorans within Genasauria, which encompasses the bulk
of the Ornithischia. The first phylogenetic definition of
the name (Table 1) employed the term Cerapoda in a
connotation equivalent to Neornithischia sensu Sereno (1999).
Indeed, Cerapoda was later defined by Weishampel (2004)
as ‘‘genasaurs more closely related to Triceratops than
to Ankylosaurus’’. The term was, however, more recently
abandoned in favour of Neornithischia (Sereno, 1997,
1998, 1999, 2005), or used in a radically different node-
based concept (Buchholz, 2002; Barrett, Butler & Knoll,
2005a; Butler et al., 2008). Originally, non-Genasauria
ornithischians included only Lesothosaurus dignosticus, from
the Early Jurassic Upper Elliot Formation, southern Africa
(Thulborn, 1970, 1972; Santa Luca, 1984; Knoll & Battail,
2001; Knoll, 2002a, b, c, 2005; Butler, 2005), and Pisanosaurus
mertii (see Section III.1). Despite its plesiomorphic postcranial
anatomy, the latter taxon has for a long time been regarded
as the sole unequivocal Triassic ornithischian, based on traits
of its partial skull and teeth (Sereno, 1991b; Butler et al., 2007;
Irmis et al., 2007b). Historically, Pisanosaurus was related to
‘‘fabrosaurids’’ (Thulborn, 1971, 1972), heterodontosaurids
(Charig & Crompton, 1974; Bonaparte, 1976; Cooper,
1985; Weishampel & Weishampel, 1983; Weishampel, 1984;
Crompton & Attridge, 1986), and ‘‘hypsilophodontids’’
(Galton, 1972, 1986; Colbert, 1981), but later accepted
as the most basal ornithischian (Novas, 1989; Weishampel
& Witmer, 1990; Sereno, 1991b; Butler, 2005; Butler et al.,
2007, 2008; but see Norman et al., 2004a). More recently, a
non-Genasauria position was also inferred for the only other
two ornithischian taxa with a Triassic record (Fig. 15A),
Heterodontosauridae (Butler et al., 2008) and Eocursor parvus
(Butler et al., 2007). This is partially based on the retention
of several features otherwise atypical for the group such
as a long hand with extensor pits on the metacarpals
and phalanges, longer penultimate phalanges, and strongly
recurved unguals with prominent flexor tubercles (Butler
et al., 2007). Heterodontosaurids are more extensively known
from Early Jurassic strata (Fig. 15B), and were traditionally
placed as the basalmost clade of Ornithopoda (Sereno, 1984,
1986; Norman, Witmer & Weishampel, 2004b), although
unorthodox alternative placements were also proposed in
the cladistic paradigm (Norman, 1984; Cooper, 1985;
Maryańska & Osmólska, 1985; Buchholz, 2002; You, Xu
& Wang, 2003; Xu et al., 2006). The group is minimally
composed of Heterodontosaurus tucki, from the Early Jurassic
Upper Elliot (Santa Luca, 1980) and Clarens (Crompton &
Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
85
Charig, 1962) formations of South Africa, and Abrictosaurus
consors, from the former unit (Thulborn, 1974). Other post-
Triassic heterodontosaurids may include Lycorhinus angustidens
(Gow, 1975), also from the Upper Elliot Formation, unnamed
forms from the Kayenta and Clarens formations (Irmis &
Knoll, 2008), as well as younger records (Norman & Barrett,
2002; Galton, 2005b; Zheng et al., 2009).
From the 1980s onwards, several putative ornithischians
(e.g. Galtonia gibbidens, Technosaurus smalli, Revueltosaurus
callenderi, R. hunti, Lucianosaurus wildi, Pekinosaurus olseni,
Tecovasaurus murryi, Protecovasaurus lucasi, Crosbysaurus harrisae)
were reported from Late Triassic assemblages of North
America (Hunt, 1989; Hunt & Lucas, 1994; Hunt et al.,
1998; Heckert, 2002, 2004) and Europe (Godefroit &
Cuny, 1997; Cuny et al., 2000; Galton, 2005b) mostly
based on isolated teeth. Yet, recent studies demonstrated
that the trustworthy Triassic record of the group is much
more restricted. Parker et al. (2005) reported the first non-
dental material of R. callenderi, showing that it represents a
pseudosuchian rather than an ornithischian. Those authors
also recognized notable convergences in the dental anatomy
of ornithischians and some non-dinosaur archosaurs, e.g.
low triangular tooth crowns with expanded base and carinae
composed of large denticles. Indeed, in a comprehensive
review of the Triassic ornithischian record, Irmis et al. (2007b)
reinterpreted most of those isolated teeth as indeterminate
archosauriforms. More recently, Ferigolo & Langer (2007)
proposed the ornithischian affinity of the purported basal
dinosauromorphs Silesaurus opolensis and Sacisaurus agudoensis
(Fig. 15A). This was partially based on the suggested
homology of the synapomorphic ornithischian predentary
bone to the edentulous tip of the lower jaw seen in both
taxa, which is formed by independent ossifications in the
latter form. Yet, that proposition was not originally backed
up by a numerical cladistic study, and further analyses
failed to recover that hypothesis of relationships (Langer &
Benton, 2006; Ezcurra, 2006; Irmis et al., 2007a; Brusatte
et al., 2008a).
Until the early eighties, most authors accepted the ‘‘fab-
rosaurids’’ as a natural group of basal ornithischians, fre-
quently depicted at the stem of either Ornithopoda or a more
inclusive group of non-thyreophoran taxa (Galton, 1978;
Norman, 1984). Yet, it became increasingly evident that ‘‘fab-
rosaurids’’ congregated a paraphyletic array of early, small-
bodied forms (Weishampel & Witmer, 1990; Sereno, 1991b).
In his phylogenetic studies, Sereno (1984, 1986, 1991b, see
also Buchholz, 2002; Xu et al., 2006) considered the archety-
pal ‘‘fabrosaurid’’ Lesothosaurus dignosticus as the sister taxon
to Genasauria. More recently, however, this view was chal-
lenged by studies that placed that form within Genasauria
(Butler et al., 2007), either as a neornithischian (Butler,
2005) or a thyreophoran (Butler et al., 2008). In the lat-
ter case, Lesothosaurus would represent the only thyreophoran
to lack the typical cortical remodeling of cranial elements
and osteoderms covering the dorsum of the body (Butler
et al., 2008). That genus was also reported from the Early
86 Max C. Langer and others

Fig. 15. Phylogenetic relationships and distribution of basal ornithischians. (A) Time-calibrated phylogeny depicting
heterodontosaurids as the most basal clade of Ornithischia, based on Butler et al. (2007, 2008); dotted lines indicate uncertain position
of Silesaurus and Sacisaurus according to Ferigolo & Langer (2007). (B) Geographic occurrences of Late Triassic (black squares) and
Early Jurassic (white squares) taxa on a Late Triassic map redrawn from Blakey (2006). (C) More conventional phylogeny depicting
heterodontosaurids as ornithopods, based on Sereno (1999); composition of Heterodontosauridae according to Butler et al. (2008).
Black silhouettes (roughly at the same scale) adapted from various sources, names applied as in Table 1. In the cladograms, node-
and stem -based taxa are respectively indicated by black circles and curved lines.

Jurassic La Quinta Formation, in western Venezuela (Rus-
sell et al., 1992), but the specimens can only be referred to
indeterminate non-cerapodan ornithischians (Barrett et al.,
2008). Other taxa previously regarded as ‘‘fabrosaurids’’
(Galton, 1978; Peng, 1997) have been considered of inde-
terminate affinity or placed within genasaurian subgroups
(Sereno, 1991b; Norman et al., 2004b, c; Butler, 2005).
Early Jurassic ornithischians other than Lesothosaurus dig-
nosticus have a less debated phylogenetic position (Fig. 15).
Scutellosaurus lawleri, from the Kayenta Formation of west-
ern USA (Colbert, 1981; Rosenbaum & Padian, 2000),
Emausaurus ernsti, from Mecklemberg, Germany (Haubold,
1991), and the genus Scelidosaurus, known from the Lower
Lias of Dorset, England (Owen, 1861, 1863; Martill, Batten
& Loydell, 2000; Norman, 2001) and possibly also from the
Kayenta Formation (Padian, 1989), are consensually con-
sidered the basalmost thyreophorans (Sereno, 1999; Butler
et al., 2008). Early Jurassic thyreophorans are otherwise only

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The origin and early evolution of dinosaurs
known from the Lufeng Formation of China (Irmis & Knoll,
2008). This includes the type specimens of Bienosaurus lufen-
gensis (Dong, 2001) and Tatisaurus oehleri (Norman, Butler &
Maidment, 2007), both of which lack autapomorphic features
and may represent nomina dubia (Irmis & Knoll, 2008). Except
for Lesothosaurus, heterodontosaurids, and thyreophorans,
Stormbergia dangershoeki, from the Upper Elliot Formation (But-
ler, 2005; but see Knoll et al., 2009), is the sole well known
Early Jurassic ornithischian. Indeed, although much richer
than during Triassic times, Early Jurassic ornithischian fau-
nas are still poorly diverse. Stormbergia dangershoeki seems to
represent the most basal Neornithischia, lacking typical fea-
tures otherwise common to the group such as an elongated
prepubic process (Butler, 2005; Butler et al., 2007, 2008).
The divergence time of Ornithischia has been traditionally
considered as Late Triassic, given the presence of Pisanosaurus
mertii and various saurischian dinosaurs in South American
deposits of that age. The phylogenetic hypothesis of Butler
et al. (2007) better fits the stratigraphic data, restricting
the Triassic record of ornithischians to non-Genasauria
taxa. Instead, both traditional (Sereno, 1999) and more
unusual (Buchholz, 2002; Xu et al., 2006) arrangements
require the existence of long ghost-lineages for Ornithopoda,
Marginocephalia, and Thyreophora. The possible position
of two South African forms at the base of both Thyreophora
and Neornithischia (Butler et al., 2008) suggests that the origin
and early diversification of ornithischians is to be found
in southwestern Gondwana (Rauhut & Lopez-Arbarello,
2008), where the Late Triassic record of the group is
concealed (Fig. 15C). On the contrary, the bulk of Early
Jurassic thyreophorans occur in North Pangea, particularly
considering the uncertain affinity of the ankylosaur reported
from the Kota Formation of India (Nath, Yadagiri & Moitra,
2002; Rauhut & Lopez-Arbarello, 2008).
The slightly more conspicuous Middle Jurassic record of
ornithischians includes the first stegosaurs and ankylosaurs
within Thyreophora and an array of basal neornithischians.
The Lower Shaximiao Formation of China (Peng et al., 2005)
has yielded stegosaurs (Maidment & Wei, 2006) and neor-
nithischians (Barrett et al., 2005a), while the former group
was recorded along with a possible pachycephalosaur from
the Balabansai Formation of Kirghizia (Averianov, Martin
& Bakirov, 2005; Averianov, Bakirov & Martin, 2007). In
Europe, Middle Jurassic ornithischians are represented by
basal ornithopods (Galton, 1980; Evans & Milner, 1994,
Kriwet, Rauhut & Gloy, 1997; Ruiz-Omeñaca, Suberbiola
& Galton, 2005), stegosaurs (Galton & Powell, 1983, Galton,
1990), and ankylosaurs (Galton, 1983a). Indeed, it was not
until the Late Jurassic and Early Cretaceous that ornithis-
chian faunas became abundant, with diverse thyreophorans,
ornithopods, ceratopsians, and pachycephalosaurs, especially
in Laurasian assemblages.
(2) Early sauropodomorph evolution
The sauropodomorph lineage includes dinosaurs with a
small skull, a distally broad humerus, a relatively short
hind limb, plus a series of other skeletal modifications
Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
87
that support their monophyly in the cladistic paradigm
(Sereno, 1999; Yates, 2003b; 2007a; Yates & Kitching, 2003;
Langer & Benton, 2006; Upchurch et al., 2007). On the
contrary, as comprehensively reviewed by Sereno (2007a;
see also Upchurch, 1997a; Galton & Upchurch, 2004),
classical studies (Huene, 1929, 1956; Romer, 1956; Colbert,
1964) frequently allocated the basal sauropodomorphs
known as ‘‘prosauropods’’ within a grade of saurischians
from which both theropods and sauropods arose. The
name Prosauropoda Huene, 1920, was coined in that
context, in reference to a group lying at the base of the
‘‘herbivorous-omnivorous’’ branch of Pachypodosauria that,
as the name implies, gave rise to sauropods. This included
Thecodontosaurus, Plateosaurus, Sellosaurus, and Anchisaurus, but
also Poekilopleuron ‘‘as a blind side stem’’ (Huene, 1920). On
the contrary, forms at some point regarded as members of
the sauropodomorph lineage such as Paleosaurus (Benton et al.,
2000) and Gresslyosaurus (Yates, 2007a) were allocated to the
theropod-related branch (Huene, 1920). As often mentioned
(Benton, 1986b), this was in part due to the mistaken
association of carnivorous teeth with other ‘‘prosauropod’’
skeletal remains (Huene, 1932; Young, 1951). Charig et al.
(1965; see also Galton, 1971, 1973) may be said to have
settled the current concept of ‘‘Prosauropoda’’, congregating
various early saurischians of the sauropodomorph lineage. As
properly put by Sereno (2007a), ‘‘prosauropods’’ represented
the first grand radiation of dinosaurs sharing minimal
morphological coherence. Basal sauropodomorphs radiated
relatively fast during Late Triassic times, becoming the
dominant terrestrial herbivores/omnivores from Norian to
Early Jurassic landscapes (Upchurch, 1997a).
Sereno (2007a) selected from the universe of basal
sauropodomorphs a subset of five taxa termed ‘‘core-
prosauropods’’, which should carry the name Prosauropoda,
if found to represent a monophylum exclusive of sauropods.
A comparable application is seen in Yates & Kitching (2003)
and Upchurch et al. (2007). However, the first phylogenetic
definition of the name was proposed under the orthodoxy of
a monophyletic ‘‘Prosauropoda’’ as to include ‘‘Thecodon-
tosauridae, Plateosauridae (Anchisauridae), Melanosauridae
[sic], and all Sauropodomorpha closer to them than to
Sauropoda’’ (Upchurch, 1997a; Sereno, 2005). Based on the
type-genera of those family rank names, this definition is not
applicable to the current framework of sauropodomorph evo-
lution, since a clade that includes Thecodontosaurus, Plateosaurus,
Anchisaurus, and Melanorosaurus also includes sauropods. Con-
sidering the criteria adopted here (Section II.2), that defi-
nition has precedence over following ones that arbitrarily
select either a single (Sereno, 1998; Galton & Upchurch,
2004) or various ‘‘prosauropods’’ (Sereno, 2007a) as inter-
nal specifiers. Indeed, the definition of Upchurch (1997a)
is to be kept and applied under an eventually revitalized
framework of ‘‘prosauropod monophyly’’, while newly pro-
posed names should be used to designate major subgroups
of Sauropodomorpha, as seen in Yates (2007a) and Smith &
Pol (2007).
88
Early evolutionary studies of basal sauropodomorphs
(Charig et al., 1965; Galton, 1971, 1976) broadly dis-
criminate three main ‘‘prosauropod’’ groups: one includ-
ing more gracile forms, the so-called ‘‘narrow-footed
prosauropods’’, termed either Thecodontosauridae or
Anchisauridae; a group of more ‘‘typical’’, Plateosaurus-related
forms; and a group of bulky quadrupeds, frequently termed
Melanorosauridae. The sauropod affinity of the latter group
was often advocated (Colbert, 1964; Cooper, 1981; Bona-
parte, 1986), hinting at ‘‘prosauropod’’ paraphyly. Indeed,
the first cladistic approach to sauropodomorph evolution
reproduced that scheme (Gauthier, 1986), allocating The-
codontosaurus antiquus and Efraasia minor at the base of the
clade, and the ‘‘broad-footed’’ forms, especially Riojasaurus
incertus, closer to Sauropoda. The succeeding cladistic studies
(Sereno, 1989, 1999; Galton, 1989; Benton et al., 2000;
Galton & Upchurch, 2004), however, failed to recover
a similar paraphyletic array of ‘‘prosauropods’’. Instead,
they pointed towards a different picture, in which most, if
not all ‘‘prosauropods’’ represented the monophyletic sis-
ter taxon to Sauropoda. This scheme is reminiscent of
pre-cladistic approaches (Charig et al., 1965; Cruickshank,
1975; Van Heerden, 1978) that partially relied on the sup-
posed irreversibility of some features of the ‘‘prosauropod’’
foot, and on the uniqueness of their hand, in order to
discard their bearing on the origin of sauropods (but see
Yates, 2003b; Sereno, 2007a). More recently, ‘‘prosauro-
pod’’ monophyly was deemed an analytical artifact derived
from poor taxon sampling, that overlooked basal and near-
sauropod sauropodomorphs (Yates, 2003b; see also Sereno,
2007a). Indeed, most recent studies (Yates & Kitching, 2003;
Pol, 2004; Smith & Pol, 2007; Yates, 2007a, b), includ-
ing those performed by previous proposers of ‘‘prosauropod
monophyly’’ (Upchurch et al., 2007; Sereno, 2007a) tend
to agree that at least some forms previously assigned to
‘‘Prosauropoda’’ are basal to the bulk of sauropodomorphs,
and that others are closely related to the sauropod radiation
(Fig. 16A). Evidently, these hypotheses are not fully congru-
ent with one another, but important common points are
seen, as outlined below.
The most recently proposed basal sauropodomorph
phylogenies (Pol, 2004; Yates, 2007a, b; Upchurch et al.,
2007) agree that the Late Triassic Saturnalia tupiniquim,
Pantydraco caducus, Thecodontosaurus antiquus, and Efraasia minor
are amongst the most basal members of the lineage, whereas
Panphagia protos may be the basal-most member (Martinez
& Alcober, 2009). Upchurch et al. (2007) also included
Mussaurus patagonicus in that basal grade, but the taxon
was given a more derived position by Pol (2004), Pol &
Powell (2005, 2007b), and Sereno (2007a) based on first-hand
examination of a more complete set of specimens. Such a
basal position was also inferred, in an admittedly tentative
fashion, to the newly described Pradhania gracilis from the
Early Jurassic Upper Dharmaram Formation, India (Kutty
et al., 2007). The relative positions of the other forms are
nearly consensual, with Saturnalia basal to Thecodontosaurus
and Pantydraco (but see Galton & Upchurch, 2004), and
Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
Max C. Langer and others
Efraasia closer to other sauropodomorphs (Fig. 16A). These
taxa represent the first radiation of the sauropodomorph
lineage, retaining various morphological traits of their
basal saurischian/basal dinosauriform precursors (Barrett
& Upchurch, 2007) such as the smaller size (adults are less
than 4 m in length) and, at least facultative, bipedality. Other
simplesiomorphies include a relatively long hand, a partially
closed acetabulum, a distal femur lacking a well-developed
extensor depression, metatarsals I and II closely appressed,
plus several other skeletal features (Yates, 2003b; Yates &
Kitching, 2003; Smith & Pol, 2007; Upchurch et al., 2007).
In addition, some of these forms possess novel herbivorous
adaptations such as a higher number of coarsely denticulated
teeth, although they might have retained an omnivorous
diet (Barrett, 2000). The geographic distribution of these
basal sauropodomorphs, along with that of slightly more
derived forms (Pol, 2004; Yates, 2007a), suggests an initial
radiation of the clade restricted to western Pangea (Fig. 16C).
Considering the node-based definition of Sauropodomorpha
and the (by typification) inclusion of Thecodontosaurus as an
internal specifier of Prosauropoda, Langer (2002) proposed
that Saturnalia should be excluded from Sauropodomorpha,
and considered instead as a taxon on its stem lineage.
The relationships of sauropodomorphs more derived than
Efraasia minor are far from consensual. In fact, several
possible arrangements recently have been proposed (see
various articles in Barrett & Batten, 2007; especially Sereno,
2007a). In most of them, however, a relatively stable set
of taxa is placed closely related to traditional sauropods
(Fig. 16A), minimally including Norian-Rhaetian forms such
as Camelotia borealis, Melanorosaurus readi, Blikanasaurus cromptoni,
and Lessemsaurus sauropoides, as well as Antetonitrus ingenipes,
which was already first described as a sauropod. This roughly
corresponds to the classical radiation of ‘‘melanorosaurids’’,
composed of usually larger (6.5-10 m), more herbivorous
‘‘prosauropods’’ (Barrett, 2000). Most of these may have
adopted a fully quadrupedal gait (Yates & Kitching, 2003),
although facultative bipedality is still suggested for several
forms (Barrett & Upchurch, 2007). These taxa share a
range of traits with eusauropods, including short and high
dorsal centra, an increased number of sacral vertebrae, a
longer manual digit I with a straighter ungual, broader non-
terminal manual phalanges, and a straighter femur, elliptical
in cross section and bearing distally displaced lesser and
fourth trochanters (Yates & Kitching, 2003; Yates, 2007a,
Upchurch et al., 2007; Pol & Powell, 2007a). Some authors
(Yates & Kitching, 2003; Smith & Pol, 2007; Yates, 2007a, b)
include all or some of these forms within Sauropoda, given the
stem-based definition of the taxon as to include forms closer
to Saltasaurus loricatus than to the archetypal ‘‘prosauropod’’
Plateosaurus engelhardti (Wilson & Sereno, 1998, Sereno, 1999;
Upchurch et al., 2007), or alternative arbitrary attempts
to mimic more traditional/current placement of forms
either within or outside the group (Yates, 2007a; Sereno,
2007a). However, Sauropoda was first phylogenetically
defined by Salgado et al. (1997) in a node-based fashion
(Table 1) that may exclude forms regularly assigned to the
The origin and early evolution of dinosaurs 89

Fig. 16. Phylogenetic relationships and distribution of basal members of the sauropodomorph lineage. (A) Time-calibrated
phylogeny depicting ‘‘core prosauropods’’ as a paraphyletic group, based on Yates (2007a, b), Smith & Pol (2007), and Martinez &
Alcober (2009); asterisk indicates alternative placement of Yunannosaurus according to Pol (2004). (B) Alternative phylogeny depicting
‘‘core prosauropod’’ monophyly, based on part of the topology proposed by Upchurch et al. (2007). (C) Geographic occurrences of
Late Triassic (black squares) and Early Jurassic (white squares) taxa on a Late Triassic map redrawn from Blakey (2006). Names
applied as in Table 1; black silhouettes (roughly at the same scale) adapted from various sources. In the cladograms, node- and stem
-based taxa are respectively indicated by black circles and curved lines.

clade such as Isanosaurus attavipachi, Gongxianosaurus shibeiensis,
Chinshakiangosaurus chuhghoensis, and Kotasaurus yamanpalliensis
(see table 2 in Upchurch et al., 2007). The latter three taxa
belong into the Early Jurassic (He, Li & Cai, 1988; Yadagiri,
2001; Upchurch et al., 2007) radiation of sauropodomorphs
that also includes some European (Wild, 1978) and North
African (Allain et al., 2004) forms. These are derived from
an already widespread set of related Triassic taxa, leading to
the well-established diversity of Mid-Late Jurassic sauropods
(Rauhut & Lopez-Arbarello, 2008).
Given such a more restrictive definition of Sauropoda,
Yates (2007a) proposed the name Massopoda (= Sauropoda
sensu Wilson & Sereno, 1998) also to encompass the
‘‘prosauropod’’ stem leading to that group, exclusive of Pla-
teosaurus engelhardti. In parallel, Sereno (2005) named a clade
composed of sauropods plus some related ‘‘prosauropods’’
as Sauropodiformes (Table 1). Apart from the taxa discussed
above, both Massopoda and Sauropodiformes probably also
include Mussaurus patagonicus (Bonaparte & Vince, 1979; Pol
& Powell, 2007b) and Jingshanosaurus xinwaensis, from the
Early Jurassic Lufeng Formation of Yunnan, China (Zhang
& Yang, 1994), a form accepted by most authors as sharing
sauropod affinities (Pol, 2004; Upchurch et al., 2007; Yates,
2007a). Additionally, newly discovered Early Jurassic forms
as Lamplughsaura dharmaramensis from the Upper Dharmaram

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90
Formation (Kutty et al., 2007), and an undescribed Argen-
tinean taxon (Pol & Garrido, 2007) may also belong to the
sauropod stem. Other Early Jurassic forms such as Yunan-
nosaurus huangi, also from the Lufeng Formation (Young,
1942), and the ‘‘prosauropod’’ from the Portland Forma-
tion of eastern North America (Anchisaurus polyzelus sensu
Yates, 2004; Ammosaurus major sensu Sereno, 2007a), have
highly controversial affinities, and two antipodal scenarios
have been presented (but see Pol, 2004). They may line
up with other sauropod-related forms (Yates, 2007a, b) or
belong to a clade of typical ‘‘prosauropods’’ (Upchurch et al.,
2007). There is limited evidence in favour of a monophyletic
group of ‘‘core prosauropods’’ (Sereno, 2007a), minimally
encompassing Plateosaurus engelhardti, Riojasaurus incertus, Mas-
sospondylus carinatus, Lufengosaurus huenei, and Coloradisaurus brevis
(Fig. 16B). Potential apomorphic traits of this clade include
modifications in the neurovascular foramina of the maxilla,
the manus (carpal II does not completely cover the proxi-
mal end of metacarpal II, metacarpal V bears an expanded
proximal end divided into two articular surfaces, first pha-
lanx of manual digit I twisted by at least 60◦ ), and foot
(metatarsal IV with expanded proximal end). On the con-
trary, Pol (2004) and Yates (2007a, b) split that group into
a successive array of three to five lineages on the stem to
Sauropodiformes sensu Sereno (2007a), where forms related
to Plateosaurus and Riojasaurus are considered more basal
(Fig. 16A). Depending on the position of these forms relative
to other sauropodomorphs is the application of names such
as Plateosauria and Anchisauria, e.g. compare Yates (2007a)
and Upchurch et al. (2007).
Either as a clade or grade, ‘‘core prosauropods’’ may
represent a biological unit, playing similar roles in the Late
Triassic-Early Jurassic ecosystems in which they occurred.
Barrett & Upchurch (2007) reviewed the palaeobiology of
these forms, highlighting some of their ecological adaptations.
These include larger size (2.5-10 m) compared to more basal
sauropodomorphs, and greater reliance on an herbivorous
diet, although facultative omnivory was not discarded. ‘‘Core
prosauropods’’ were also capable of bipedal locomotion
(Cooper, 1981; Christian & Preuschoft, 1996; Bonnan
& Senter, 2007), although larger forms were probably
obligatory quadrupeds. This is the case for Riojasaurus incertus,
previously connected to the origin of sauropods (Bonaparte,
1972; Gauthier, 1986). As a whole, that clade/grade
congregates a relatively high diversity of Late Triassic forms,
including Ruehleia bedheimensis and the species of Plateosaurus
in Europe/Greenland, Riojasaurus incertus and Unaysaurus
tolentinoi in South America, as well as Eucnemesaurus fortis
and Plateosauravus cullingworthi in South Africa. Other ‘‘core-
prosauropods’’ may fit into Massospondylidae, a clade that
includes the Triassic Coloradisaurus brevis (Yates & Kitching,
2003), the Early Jurassic Glacialisaurus hammeri (Smith & Pol,
2007) from the Hanson Formation, Antarctica, and a possible
set of Chinese forms (see Pol, 2004), minimally including
Lufengosaurus huenei from the Lufeng Formation (Barrett,
Upchurch & Xiao-Lin, 2005b). Its type genus Massospondylus
(M. carinatus) is known from the Upper Elliot Formation
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Max C. Langer and others
(Cooper, 1981; Gow, Kitching & Raath, 1990; Sues et al.,
2004, Reisz et al., 2005) and other stratigraphic units in
southern Africa (Cooper, 1981; Galton & Upchurch, 2004),
but not in North America (Attridge, Crompton & Jenkins,
1985; Sues et al., 2004). More recently, its sister taxon,
Adeopapposaurus mognai, was described from the Lower Jurassic
Cañon del Colorado Formation, Argentina (Martinez, 2009).
(3) Early theropod evolution
The name Theropoda was coined by Marsh (1881) as
a new suborder of carnivorous dinosaurs, but its status
as a ‘‘natural group’’ was rejected for the first half of
the last century. At the time, gracile members of the
group, the so-called ‘‘coelurosaurs’’, were considered, along
with ‘‘prosauropods’’, the ‘‘basal stock’’ from which both
sauropods and derived theropods evolved (Huene, 1914,
1920, 1932; Romer, 1956). Theropod monophyly was hinted
at by Matthew & Brown (1922), firmly established by Col-
bert (1964), in an arrangement widely accepted since (Charig
et al., 1965; Romer, 1966; Colbert & Russell, 1969; Ostrom,
1978; Steel, 1970; Bakker & Galton, 1974), and corrobo-
rated by pioneering phylogenetic studies (Thulborn, 1984;
Gauthier, 1986; Novas, 1992b; Holtz, 1994; Sereno, 1999).
Indeed, taxa consensually assigned to the group share a series
of typical traits, e.g. promaxillary foramen; well-developed
pneumatization in cervical and cranial trunk vertebrae;
manus with reduced metacarpal I, slender metacarpal III,
and reduced/absent digits IV and V; ilium with promi-
nent supracetabular crest and preacetabular ala; tibia with
marked cnemial and fibular crests; transversely compressed
calcaneum; foot with reduced outer digits (Rauhut, 2003;
Ezcurra & Cuny, 2007; Ezcurra & Novas, 2007a).
As discussed in Section III.2, the most contentious
aspect of early theropod evolution is the possible nesting
of various Triassic forms within the group (Fig. 9). This
is particularly the case fot Eoraptor lunensis (Sereno et al.,
1993) and herrerasaurs (Sereno & Novas, 1992), but also
for other taxa such as Guaibasaurus candelariensis (Langer et al.,
2007a), Agnosphitys cromhallensis, and Chindesaurus bryansmalli
(Yates, 2007a). Indeed, these forms apart, the Norian
coelophysoid ‘‘Camposaurus arizonensis’’ represents the oldest
theropod (Hunt et al., 1998; Nesbitt et al., 2007), which would
make Theropoda the only major dinosaur lineage lacking a
well-defined Ischigualastian record. Moreover, based on the
current knowledge of theropod diversity, and not considering
the above-mentioned taxa as members of the group, the
stem-based Theropoda would be as inclusive as the node-
based Neotheropoda (Table 1). This name was first used
by Bakker (1986) to combine theropods more derived than
‘‘podokesaurids’’, but phylogenetically defined by Sereno
(1998) as a more inclusive group. Yet, Neotheropoda remains
a useful name under alternative arrangements (Fig. 9) and/or
if new forms are found to belong to its stem (see Nesbitt &
Chatterjee, 2008; Martinez et al., 2008). Although a less
inclusive Neotheropoda (Padian et al., 1999; Wilson et al.,
2003) seems more useful in the current orthodoxy, and more
properly translates the original meaning of the name (Bakker,
The origin and early evolution of dinosaurs
1986), the definition proposed by Sereno (1998) has historical
‘‘priority’’.
Another controversial aspect of early theropod evolution is
the possible monophyly of the oldest neotheropods, grouped
within Ceratosauria and/or Coelophysoidea. Indeed, the first
cladistic analyses of theropod relationships identified two
main neotheropod lineages, Ceratosauria and Tetanurae
(Gauthier, 1986; Rowe, 1989; Rowe & Gauthier, 1990).
In turn, Ceratosauria was divided into two branches, the
Late Triassic-Early Jurassic Coelophysoidea (i.e. Coelophysis,
Dilophosaurus, and their kin) and the Jurassic-Cretaceous
Neoceratosauria, including Ceratosaurus and abelisauroids
(Novas, 1992a; Sereno, 1997, 1999; Holtz, 2000; Coria &
Salgado, 2000). This arrangement was accepted during most
of the 1990s, but the vast majority of more recent studies
consider neoceratosaurs more closely related to tetanurans
than to coelophysoids, challenging the monophyly of
the traditional Ceratosauria (Carrano, Sampson & Forster,
2002; Carrano, Hutchinson & Sampson, 2005; Rauhut,
2003; Sereno, Wilson & Conrad, 2004; Yates, 2005; Ezcurra,
2006; Ezcurra & Novas, 2007a; Ezcurra & Cuny, 2007; Smith
et al., 2007; Carrano & Sampson, 2008; but see Tykoski &
Rowe, 2004; Tykoski, 2005; Allain et al., 2007). In fact,
Ceratosauria was node-based defined by Rowe & Gauthier
(1990) as ‘‘including Ceratosaurus nasicornis, Dilophosaurus
wetherilli, Liliensternus liliensterni, Coelophysis bauri, Syntarsus
rhodesiensis, Syntarsus kayentakatae, Segisaurus halli, Sarcosaurus
woodi, and all other taxa stemming from their most recent
common ancestor’’, based on a phylogenetic framework
in which these forms compose a monophylum exclusive
of tetanurans. On the contrary, in the current orthodoxy
(Fig. 17), Ceratosauria would point to the same node as
Neotheropoda, circumscribing a much more inclusive group,
and is not employed here. Instead, the term Averostra
Paul, 2002, as phylogenetically defined by Ezcurra & Cuny
(2007), designates the clade composed of Tetanurae plus
Neoceratosauria (Table 1).
Early phylogenetic studies grouped all Triassic and
Early Jurassic neotheropods within the Coelophysoidea
clade. This included Dilophosaurus wetherilli, as the sister
taxon to Liliensternus liliensterni plus Coelophysidae (Rowe
& Gauthier, 1990; Holtz, 1994). The latter group is
minimally composed of Late Triassic (Colbert, 1989; Rauhut
& Hungerbühler, 2000) and Early Jurassic (Raath, 1969;
Rowe, 1989) Coelophysis-‘‘Syntarsus’’-related forms (Paul, 1988;
Bristowe & Raath, 2004), but may also include Segisaurus
halli and Procompsognathus triassicus (Sereno, 1997, 1999;
Tykoski & Rowe, 2004; Knoll, 2008). With the addition
of newly described forms, such as Zupaysaurus rougieri and
Lophostropheus airelensis, various cladisitic analyses (Tykoski &
Rowe, 2004; Carrano et al., 2005; Tykoski, 2005; Ezcurra,
2006; Ezcurra & Novas, 2007a; Ezcurra & Cuny, 2007)
also recovered a monophyletic Coelophysoidea sensu lato
(Fig. 17B). As such, the group can be diagnosed by several
craniomandibular features such as a flexible articulation
between premaxilla and maxilla marked by a prominent
subnarial diastema, an expanded rostral end of the dentary,
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91
reduced serrations on premaxillary teeth, and enlarged (fang-
like) teeth in the rostral portion of the dentary, as well as
by a peculiar pattern of femoral dimorphism (Tykoski &
Rowe, 2004; Ezcurra & Novas, 2007a, Ezcurra & Cuny,
2007). However, the monophyletic status of that group
was questioned by Rauhut (2003), who found Dilophosaurus
more closely related to tetanurans and neoceratosaurs than
to coelophysids. Along with other forms, those taxa share
cranial details such as a lacrimal fenestra, a dorsoventrally
elongated orbit, and a reduced tooth count occupying
a shorted portion of the maxilla, modifications in the
retroarticular process of the lower jaw, and a higher astragalar
ascending process (Smith et al., 2007). A similar hypothesis of
relationship was advocated by Yates (2005), which considered
Dracovenator regenti as closely related to Dilophosaurus wetherilli.
Smith et al. (2007) also found other Early Jurassic taxa
such as Cryolophosaurus ellioti and ‘‘Dilophosaurus’’ sinensis
as members of such a ‘‘Dilophosaurus clade’’ (Fig. 17A),
partially characterized by the presence and/or differential
configuration of the dorsal crests of the skull. Yates (2005)
also assigned Zupaysaurus to that clade, but the phylogenetic
position of that taxon is highly controversial within non-
averostran neotheropods (Carrano et al., 2005; Smith et al.,
2007; Ezcurra & Novas, 2007a). Clearly, as stem-based-
defined by Sereno (1998), the inclusivity of Coelophysoidea
is dependent on the adopted evolutionary framework. It
may include only small to medium-sized forms similar
to Coelophysis-‘‘Syntarsus’’ (Fig. 17A), or also congregate an
Early Jurassic radiation of large-bodied Dilophosaurus-like
taxa (Fig. 17B).
Neotheropods experienced a rapid diversification during
the Norian-Rhaetian and Early Jurassic, achieving a broad
distribution over west Pangea (Fig. 17C). This early radiation
was mostly represented by sensu lato ‘‘coelophysoids’’, i.e.
non-averostran neotheropods. Their Norian representatives
include small to medium-sized forms reported from western
USA (Colbert, 1989; Carpenter, 1997; Nesbitt et al., 2007)
and Europe (Sereno & Wild, 1992; Rauhut & Hungerbühler,
2000; Allen, 2004), most of which are also sensu sticto
Coelophysoidea (Fig. 17B), and perhaps also larger forms
such as Zupaysaurus rougieri (Arcucci & Coria, 2003).
Close to the Triassic-Jurassic boundary, theropod remains
become scarce and basically only Lophostropheus airelensis
is known (Ezcurra & Cuny, 2007; but see Dzik et al.,
2008). Tetanurans previously have been reported from Late
Triassic outcrops, but these records are not conclusive. The
supposed bird Protoavis texensis (Chatterjee, 1991) has been
recently reinterpreted as a chimaera (Nesbitt et al., 2007),
with some elements of ‘‘coelophysoid’’ affinities, but not
of tetanuran nature. Otherwise, ‘‘Zanclodon’’ cambrensis was
assigned to Tetanurae (Holtz, Molnar & Currie, 2004),
but the available material does not differ from those of
non-averostran theropods.
During the Early Jurassic (Fig. 17C), sensu sticto coelo-
physoids continue to be well represented in western USA,
including Segisaurus halli from the Navajo Sandstone (Camp,
1936; Carrano et al., 2005) and ‘‘Syntarsus’’ kayentakatae from
92 Max C. Langer and others

Fig. 17. Phylogenetic relationships and distribution of basal theropods. (A) Time calibrated phylogeny depicting ‘‘coelophysoids’’
as a paraphyletic group, based on Smith et al. (2007); position of Lophostropheus according to Ezcurra & Cuny (2007); position of
Procompsognathus and relations within Coelophysidae according to Tykoski & Rowe (2004); asterisk indicates alternative placement
of Zupaysaurus as sister taxon to Dracovenator, according to Yates (2005); dotted lines indicate uncertain position of herrerasaurids
and Eoraptor according to Sereno (1999) and Guaibasaurus according to Langer & Benton (2006). (B) Alternative phylogeny depicting
‘‘coelophysoid’’ monophyly, based on Ezcurra & Cuny (2007) and Ezcurra & Novas (2007a); asterisk indicates alternative placement
of Zupaysaurus according to Carrano et al. (2005). (C) Geographic occurrences of Late Triassic (black squares) and Early Jurassic
(white squares) taxa on a Late Triassic map redrawn from Blakey (2006). Names applied as in Table 1; black silhouettes (roughly at
the same scale) adapted from various sources. In the cladograms, node- and stem -based taxa are respectively indicated by black
circles and curved lines.

the Kayenta Formation (Rowe, 1989), both in Arizona. In
addition, ‘‘Coelophysis’’ rhodesiensis is known from the Upper
Elliot Formation of South Africa (Raath, 1980) and especially
from the Forest Sandstone of Zimbabwe (Raath, 1969). Fur-
ther Jurassic records of ‘‘coelophysoids’’ are known from
China (Irmis, 2004), Mexico (Munter & Clark, 2006), and
possibly Europe (Andrews, 1921; Carrano & Sampson, 2004).
Larger forms attributed or not to the Dilophosaurus-clade
also retain a broad distribution. These include Dilophosaurus
wetherilli from the Kayenta Formation (Welles, 1984), Dra-
covenator regenti from the Upper Elliot Formation (Yates,
2005), Cryolophosaurus ellioti from the Hanson Formation
of the Transantarctic Mountains (Hammer & Hickerson,
1994; Smith et al., 2007), and ‘‘Dilophosaurus’’ sinensis from

Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
The origin and early evolution of dinosaurs
the Lufeng Formation of China (Hu, 1993). In addition,
the oldest averostrans are known from Early Jurassic assem-
blages. This includes Berberosaurus liassicus, a neoceratosaur
from Morocco (Allain et al. 2007; but see Xu et al., 2009),
and the very dubious record of a therizinosauroid jaw in the
Lufeng Formation (Zhao & Xu, 1998; Xu, Zhao & Clark,
2001; Rauhut, 2003). In any case, as sister taxon to Neo-
ceratosauria, a tetanuran ghost lineage might be inferred for
the latest Early Jurassic (Fig. 17A). Further Early Jurassic
theropods were recorded from the La Quinta Formation of
Venezuela (Moody, 1997) based on teeth that can not be
allocated in a less inclusive clade.
Middle Jurassic dinosaur-bearing assemblages are rare
(Rauhut & Lopez-Arbarello, 2008), but the available data
show that the composition of theropod faunas changed
drastically relative to those of Late Triassic and Early
Jurassic age. ‘‘Coelophysoids’’ disappear completely from the
fossil record, and tetanurans became the dominant forms.
Apart from indeterminate theropod remains from North
Africa (Monbaron, Russell & Taquet, 1999) and Madagascar
(Flynn et al., 2006), Middle Jurassic forms include a probable
basal neoceratosaur from Australia (Long & Molnar, 1998;
Rauhut, 2005a), and basal tetanurans form Argentina
(Rauhut, 2005a), Europe, and China (Holtz et al., 2004;
Smith et al., 2007). On the other hand, neoceratosaurs are
better known from Late Jurassic and especially Cretaceous
deposits (Carrano & Sampson, 2008; Xu et al., 2009).
VI. CONCLUSIONS
(1) The oldest unequivocal records of Dinosauria are of
Late Triassic age (approximately 230 Mya). These were
unearthed from rocks accumulated over extensional rift
basins in Argentina, Brazil, Zimbabwe, and India. The
better known early dinosaurs are Herrerasaurus ischigualastensis,
Pisanosaurus mertii, Eoraptor lunensis, and Panphagia protos from
the Ischigualasto Formation, northwestern Argentina, and
Staurikosaurus pricei and Saturnalia tupiniquim from the Santa
Maria Formation, south Brazil. Other dinosaur records of
equivalent age are either more fragmentary or of dubious
affinities, hinting at a south Pangea origin of the group. No
uncontroversial dinosaur body fossils are known from older
strata, but a possible Middle Triassic origin of the lineage
may be inferred from both the footprint record and its sister
group relation to Ladinian basal dinosauromorphs.
(2) Dinosauria is by definition a monophyletic group that,
in the present orthodoxy, combines saurischians and ornithis-
chians to the exclusion of other major archosaur groups such
as pterosaurs, phytosaurs, and crocodylomorphs. The first
phylogenetic definition to fit the current understanding of
Dinosauria as a node-based taxon solely composed of mutu-
ally exclusive Saurischia and Ornithischia was given as ‘‘all
descendants of the most recent common ancestor of birds
and Triceratops’’. This definition should be followed until
more specific provisions are given by the PhyloCode.
Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
93
(3) Dinosaurs are nested within the bird-line of archosaurs
along with pterosaurs (possibly), Scleromochlus taylori, and basal
Dinosauromorpha, the phylogeny of which is in state of
flux. It includes the archetypal Marasuchus lilloensis, a diver-
sity of more basal forms such as Lagerpeton and Dromomeron,
as well as silesaurids: a possibly monophyletic group that
combines Mid-Late Triassic basal dinosauromorphs that
may represent sister taxa to Dinosauria. Recent cladistic
analyses of basal dinosaur relationships agree in various
key points: (1) Pisanosaurus mertii is a basal ornithischian;
(2) Herrerasaurus ischigualastensis and Staurikosaurus pricei belong
in a monophyletic Herrerasauridae; (3) Guaibasaurus candelar-
iensis, Eoraptor lunensis, and herrerasaurids are saurischians;
(4) Saurischia includes two main groups, Sauropodomor-
pha and Theropoda; and (5) Saturnalia tupiniquim is a basal
member of the sauropodomorph lineage, a position also
inferred for the recently described Panphagia protos. On the
contrary, several aspects of basal dinosaur phylogeny remain
controversial, including the position of silesaurids as basal
ornithischians or non-dinosaur dinosauromorphs; the posi-
tion of herrerasaurids, Eoraptor, and Guaibasaurus as basal
theropods or basal saurischians; and the affinity and/or valid-
ity of various fragmentary taxa such as Agnosphitys cromhallensis,
Alwalkeria maleriensis, Chindesaurus bryansmalli, Saltopus elginensis,
Spondylosoma absconditum, and Teyuwasu barberenai.
(4) The identification of dinosaur apomorphies is
hampered by the incompleteness of the skeletal remains
attributed to most basal dinosauromorphs, the skull and fore
limb of which are particularly poorly known. Nonetheless,
to the exclusion of silesaurids, Dinosauria can be diagnosed
by a suite of derived traits, namely: foramen-sized post-
temporal fenestra; epipophyses on cranial cervical vertebra;
long deltopectoral crest; open acetabulum; arched dorsal
iliac margin; femoral head inturned and distinctly offset from
the shaft; asymmetrical fourth trochanter; astragalus with
acute anteromedial corner, broad ascending process, and
reduced fibular articulation; and proximally flat lateral distal
tarsal. On the contrary, long-standing dinosaur apomorphies
such as the absence of a postfrontal, the presence of more
than two sacral vertebrae, reduced manual digits IV and
V, modified ‘‘lesser trochanter’’, and metatarsals II and IV
subequal in length do not unambiguously diagnose the group.
The prevalence of dinosaur diagnostic traits related to the
pelvic girdle and limb may reflect the better preservation
of these structures in their sister taxa, but may also suggest
that these anatomical parts suffered most of the changes seen
in the early dinosaur skeleton. Some of these traits can be
related to the acquisition of an erect bipedal gait, which has
traditionally been suggested to represent a key adaptation
that allowed, or even promoted, dinosaur radiation in the
Late Triassic.
(5) Contrary to the classical ‘‘competitive’’ model,
dinosaurs did not gradually replace other terrestrial tetrapods
over the Late Triassic. Yet, opportunistic and competitive
scenarios are not mutually exclusive, and species interaction
may have played a partial role in the rise of dinosaurs,
which can be said to have consisted of three landmark
94
moments, separated by controversial (Carnian-Norian,
Triassic-Jurassic) extinction events. The Carnian early
diversification did not occur in an empty ecospace but despite
the abundance and diversity of contemporary tetrapods.
The Norian increase in dominance might be connected to
climatic/floristic changes and to the extinction of herbivorous
forms such as rhynchosaurs, but the timing of these events
needs further investigation. The subtle Jurassic diversification
seems to have occurred in the aftermath of an extinction
event, and might be an example of opportunistic radiation
into released ecospace, with the origin of neornithischians
and thyreophorans, the heterodontosaurid diversity peak,
the rise of the Dilophosaurus-clade, and further acquisition of
typical graviportal traits among sauropods.
(6) It is traditionally believed that dinosaurs arose from
bipedal and carnivorous forms, but evidence gathered
from newly discovered basal dinosauromorphs indicate
that quadrupedalism and omnivory/herbivory can not be
discarded as possible ancestral traits of the group. At the
moment, however, most evidence points towards a fully
bipedal origin of dinosaurs. On the contrary, depending
on the accepted hypothesis of relationships, the ancestral
dinosaur diet can be reconstructed as either carnivorous
or omnivorous. In any case, the plesiomorphic tooth
morphology of dinosaurs does not strictly compare to the
typical carnivorous or omnivorous/herbivorous patterns of
more derived members of the group. Indeed, each major
dinosaur group seems to have independently acquired its
typical set of dental traits.
(7) The phylogenetic relationships of the basal members
of each major dinosaur group have recently been reeval-
uated in the light of new evidence. Among ornithischians,
unorthodox placements inside and outside Genasauria were
proposed for Lesothosaurus diagnosticus and Heterodontosauri-
dae, respectively. Within saurischians, both ‘‘Prosauropoda’’
and ‘‘Ceratosauria’’ were regarded as paraphyletic in their
broader understanding. Yet, ‘‘core prosauropods’’ and coelo-
physoids may still represent smaller clades at the base of
Sauropodomorpha and Theropoda, respectively.
(8) Whereas the oldest dinosaurs were geographically
restricted to south Pangea, including rare ornithischians
and more abundant basal members of the saurischian
lineage, the group achieved a nearly global distribution
by Norian/Rhaetian times, especially with the radiation
of saurischian groups such as ‘‘prosauropods’’ and coelo-
physoids. This suggests an ‘‘out of south Pangea’’ model of
dinosaur radiation, but no model is better than the evidence
upon which it stands. In this case, the evidence is restricted
to a handful of fossils from scattered areas around the world,
and more prospection work is needed in order to build a more
reliable scenario of dinosaur origins and basal radiation.
VII. ACKNOWLEDGEMENTS
We thank the following for permission to examine specimens
mentioned in this paper and help during this work: Angela
Biological Reviews 85 (2010) 55–110 © 2009 The Authors. Journal compilation © 2009 Cambridge Philosophical Society
Max C. Langer and others
Milner and Sandra Chapman (BMNH); Rainer Schoch and
Rupert Wild (SMNS); Ricardo Martinez and Oscar Alcober
(UNSJ); Dave Unwin (MB); Pat Holroyd, Randy Irmis,
and Kevin Padian (UCMP); Michael Maisch (GPIT); Jaime
Powell and Judith Babot (PVL); Mike Raath (QVM); Jerzy
Dzik and Tomasz Sulej (ZPAL); Maria Claudia Malabarba
(MCP); Jorge Ferigolo and Ana Maria Ribeiro (FZB/RS);
and Emilio Vaccari (PULR). Aspects of this work were funded
by grants from the Brazilian agencies CNPq, CAPES, and
FAPESP (to M.C.L. and J.S.B.), Samuel Welles Fund–UC
Berkeley (to M.D.E.), Agencia Nacional de Promoción
Cientı́fica y Técnica (to F.E.N.), and Santander Bank (to
J.S.B.). Comments by Mike Benton and an anonymous
reviewer were much appreciated. This is contribution No.
15 of Laboratório de Paleontologia, FFCLRP-USP.
VIII. APPENDIX 1. INSTITUTIONAL
ABBREVIATIONS
BMNH, Natural History Museum, London, UK; GPIT,
Institut für Geologie und Paläontologie, Tübingen, Ger-
many; MB, Humboldt Museum für Naturkunde, Berlin,
Germany; MCP, Museu de Ciências e Tecnologia, PUCRS,
Porto Alegre, Brazil; PULR, Museo de Ciencias Naturales,
Universidad Nacional de La Rioja, La Rioja, Argentina;
PVL, Fundación ‘‘Miguel Lillo’’, San Miguel de Tucumán,
Argentina; PVSJ, Museo de Ciencias Naturales, Universidad
Nacional de San Juan, San Juan Argentina; QVM, National
Museum of Natural History, Harare, Zimbabwe; SMNS,
Staatliches Museum für Naturkunde, Stuttgart, Germany;
UCMP, University of California Museum of Paleontology,
Berkeley, California, USA; ZPAL, Institute of Paleobiology
of the Polish Academy of Science, Warsaw, Poland.
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